Chinyorwa ichi chikamu chemusoro wenyaya yekutsvagisa unoti "Tekinoroji dzepamusoro dzekugadzirisa zvinhu zvipenyu uye maitiro ekugadzirisa zvinhu zvakagadzirwa nemakemikari emuchadenga (SOC). Ona zvinyorwa zvese gumi nemana
Mapuroteni ane huremu hwakaderera epolycyclic aromatic hydrocarbons (PAHs) akadai senaphthalene uye naphthalenes dzakatsiviwa (methylnaphthalene, naphthoic acid, 1-naphthyl-N-methylcarbamate, nezvimwewo) anoshandiswa zvakanyanya mumaindasitiri akasiyana-siyana uye ane chepfu, mutagenic uye/kana carcinogenic kune zvipenyu. Aya macompounds ekugadzira (SOCs) kana xenobiotics anoonekwa seanonyanya kusvibisa uye anotyisidzira zvakanyanya nharaunda yepasi rose nehutano hweveruzhinji. Kusimba kwemabasa evanhu (semuenzaniso gasification yemarasha, kuchenesa mafuta, kuburitsa kwemotokari uye mashandisirwo ekurima) kunosarudza huwandu, mugumo uye kutakurwa kwemakemikari aya aripo uye anogara aripo. Kuwedzera kune nzira dzekurapa/kubvisa makemikari nemakemikari, matekinoroji egirini uye ane hushamwari kune nharaunda akadai se bioremediation, ayo anoshandisa tupukanana tunogona kukanganisa zvachose maPOC kana kuashandura kuita zvigadzirwa zvisina chepfu, abuda senzira yakachengeteka, inodhura uye inovimbisa. Mhando dzakasiyana dzebhakitiriya dziri mu phyla Proteobacteria (Pseudomonas, Pseudomonas, Comamonas, Burkholderia, uye Neosphingobacterium), Firmicutes (Bacillus naPaenibacillus), uye Actinobacteria (Rhodococcus naArthrobacter) mu ivhu microbiota dzakaratidza kugona kushatisa makemikari akasiyana-siyana e organic. Zvidzidzo zve metabolism, genomics, uye metagenomic analysis zvinotibatsira kunzwisisa kuomarara kwe catabolic uye kusiyana kuripo mumhando idzi dzehupenyu dziri nyore, izvo zvinogona kushandiswa zvakare kuti biodegradation ibudirire. Kuvapo kwenguva refu kwePAHs kwakaguma nekubuda kwemaitiro matsva ekuora kuburikidza nekutamiswa kwemajini akatenderera uchishandisa zvinhu zvemajini zvakaita seplasmids, transposons, bacteriophages, genomic islands, uye integrative conjugative elements. Systems biology uye genetic engineering ye specific isolates kana model communities (consortia) zvinogona kugonesa bioremediation yakakwana, inokurumidza uye inoshanda yePAHs idzi kuburikidza ne synergistic effects. Muongororo iyi, tinotarisa panzira dzakasiyana dzekugadzirisa muviri nekusiyana-siyana, kuumbwa kwemajini nekusiyana-siyana, uye mhinduro dzemasero/kugadziriswa kwenaphthalene uye mabhakitiriya anoparadza naphthalene panzvimbo peimwe. Izvi zvichapa ruzivo rwezvakatipoteredza rwekushandiswa mumunda uye kugadzirisa kusvuta kuti zvinhu zvirambe zvichishanda zvakanaka.
Kukurumidza kukura kwemaindasitiri (makemikari epetrochemicals, kurima, mishonga, madhayi emachira, zvizoro, nezvimwewo) kwakabatsira mukubudirira kwehupfumi hwenyika uye kuvandudza mararamiro. Kubudirira uku kwakakonzera kugadzirwa kwehuwandu hwakawanda hwemakemikari ekugadzira (SOCs), anoshandiswa kugadzira zvigadzirwa zvakasiyana-siyana. Aya makemikari ekunze kana kuti maSOC anosanganisira polycyclic aromatic hydrocarbons (PAHs), mishonga yezvipembenene, mishonga yekuuraya zvipembenene, mapurasitiki, madhayi, mishonga yemishonga, organophosphates, zvinodzivisa moto, zvinonyungudutsa organic solvents, nezvimwewo. Anoburitswa mumhepo, mumvura uye pasirese uko ane mhedzisiro yakawanda, zvichikonzera mhedzisiro yakaipa pamhando dzakasiyana dzezvinhu zvipenyu kuburikidza nekuchinja kwehunhu hwefizikisi uye chimiro chenharaunda (Petrie et al., 2015; Bernhardt et al., 2017; Sarkar et al., 2020). Zvinhu zvakawanda zvinosvibisa zvinonhuhwirira zvine simba uye zvinoparadza pazvipenyu zvakawanda zvisina kukuvadzwa/nzvimbo dzinopisa zvipenyu (semuenzaniso matombo ekorari, Arctic/Antarctic ice sheets, madhamu marefu emakomo, madhaka egungwa rakadzika, nezvimwewo) (Jones 2010; Beyer et al. 2020; Nordborg et al. 2020). Zvidzidzo zve geomicrobiological zvechangobva kuitwa zvakaratidza kuti kuiswa kwezvinhu zvakasikwa (semuenzaniso zvinosvibisa zvinonhuhwirira) nezvinhu zvinobva mazviri pamusoro pezvivakwa zvekugadzira (nzvimbo dzakavakwa) (semuenzaniso nzvimbo dzenhaka dzetsika nematombo akagadzirwa negranite, matombo, huni nesimbi) zvinokurumidzisa kuora kwazvo (Gadd 2017; Liu et al. 2018). Mabasa evanhu anogona kuwedzera uye kuwedzera kuipa kwekuparadzwa kwezvinhu zvipenyu nezvivakwa kuburikidza nekusvibiswa kwemhepo nekushanduka kwemamiriro ekunze (Liu et al. 2020). Izvi zvinosvibisa zvipenyu zvinosangana nemhepo mumhepo uye zvinogara pachivako, zvichikonzera kuora kwemuviri nemakemikari kwezvinhu. Kuora kwezvipenyu kunozivikanwa zvakanyanya sekuchinja kusingadiwe pakuonekwa uye hunhu hwezvinhu zvinokonzerwa nezvipenyu zvinokanganisa kuchengetedza kwazvo (Pochon naJaton, 1967). Kufambira mberi kwekuita kwehutachiona (metabolism) hwezvinhu izvi kunogona kuderedza kusimba kwechimiro, kushanda zvakanaka kwekuchengetedza uye kukosha kwetsika (Gadd, 2017; Liu et al., 2018). Kune rumwe rutivi, mune dzimwe nguva, kugadziriswa kwehutachiona uye kupindura kune izvi zvivakwa kwakaonekwa kuti kunobatsira sezvo zvichigadzira biofilms nezvimwe zvidimbu zvinodzivirira zvinoderedza mwero wekuora/kuora (Martino, 2016). Nokudaro, kuvandudzwa kwenzira dzinoshanda dzekuchengetedza kwenguva refu dzematombo, simbi nematanda zvinoda kunzwisisa kwakakwana maitiro akakosha anobatanidzwa mukuita uku. Zvichienzaniswa nemaitiro echisikigo (maitiro e geological, moto wemasango, kuputika kwemakomo, maitiro emiti nemabhakitiriya), mabasa evanhu anokonzera kuburitswa kwehuwandu hwakawanda hwe polycyclic aromatic hydrocarbons (PAHs) nedzimwe organic carbon (OC) mu ecosystems. MaPAH mazhinji anoshandiswa mukurima (mishonga yezvipembenene nemishonga yezvipembenene yakaita seDDT, atrazine, carbaryl, pentachlorophenol, nezvimwewo), maindasitiri (mafuta asina kuchena, tsvina yemafuta, mapurasitiki anobva mupetroleum, maPCB, mapurasitiki, sipo, mishonga yekuuraya utachiona, fumigants, zvinonhuhwirira uye zvinochengetedza), zvigadzirwa zvekuchengetedza muviri (zvinodzivirira zuva, mishonga yekuuraya utachiona, zvinodzinga zvipembenene uye polycyclic musks) uye zvombo (zvinoputika zvakaita se2,4,6-TNT) zvinogona kukonzera xenobiotics zvinogona kukanganisa hutano hwepasi (Srogi, 2007; Vamsee-Krishna naPhale, 2008; Petrie et al., 2015). Rondedzero iyi inogona kuwedzerwa kusanganisira mishonga inobva mupetroleum (mafuta emafuta, mafuta emafuta, asphaltenes), bioplastics ine huremu hwakawanda, uye ionic liquids (Amde et al., 2015). Tafura 1 inonyora mhando dzakasiyana dzezvinonhuhwirira zvinosvibisa uye mashandisirwo azvo mumaindasitiri akasiyana-siyana. Mumakore achangopfuura, kuburitswa kwezvinhu zvinoshanduka kubva mumhepo, pamwe necarbon dioxide nemamwe magasi egreenhouse, kwakatanga kuwedzera (Dvorak et al., 2017). Zvisinei, kukanganisa kwevanhu kunopfuura zvakanyanya kwezvisikwa. Pamusoro pezvo, takaona kuti maSOC akati wandei anoramba aripo munzvimbo dzakawanda dzezvakatipoteredza uye akaonekwa seanobuda achisvibisa zvinhu zvine mhedzisiro yakaipa pazvipenyu (Mufananidzo 1). Masangano ezvakatipoteredza akadai seUnited States Environmental Protection Agency (USEPA) akaisa akawanda ezvinosvibisa izvi pane zvavanokoshesa nekuda kwehunhu hwavo hunokonzera cytotoxic, genotoxic, mutagenic, uye carcinogenic. Nokudaro, mitemo yakasimba yekurasa marara uye nzira dzinoshanda dzekurapa/kubvisa marara kubva munzvimbo dzakasvibiswa dzinodiwa. Nzira dzakasiyana-siyana dzekurapa panyama nemakemikari dzakadai se pyrolysis, oxidative thermal treatment, air aeration, landfilling, incineration, etc. hadzina basa uye dzinodhura uye dzinogadzira zvinoparadza, zvine chepfu uye zvakaoma kugadzirisa. Nekuwedzera kweruzivo rwezvakatipoteredza pasi rose, tupukanana tunokwanisa kuparadza tsvina iyi nezvinobuda mairi (zvakaita se halogenated, nitro, alkyl uye/kana methyl) turi kukwezva kutariswa kuri kuwedzera (Fennell et al., 2004; Haritash naKaushik, 2009; Phale et al., 2020; Sarkar et al., 2020; Schwanemann et al., 2020). Kushandiswa kwetupukanana utwu chete kana mumarudzi akasiyana-siyana (makoloni) kubvisa tsvina inonhuhwirira kune zvakanakira maererano nekuchengetedzeka kwezvakatipoteredza, mutengo, kushanda zvakanaka, kushanda zvakanaka, uye kugara kwenguva refu. Vaongorori vari kuongororawo kubatanidzwa kwemaitiro etupukanana nenzira dze electrochemical redox, dzinoti bioelectrochemical systems (BES), se tekinoroji inovimbisa yekurapa/kubvisa tsvina (Huang et al., 2011). Tekinoroji yeBES yakwezva kutariswa kuri kuwedzera nekuda kwekushanda kwayo kwepamusoro, mutengo wakaderera, kuchengetedzeka kwezvakatipoteredza, kushanda kwetembiricha yemukamuri, zvinhu zvinoenderana nehupenyu, uye kugona kudzoreredza zvigadzirwa zvinokosha (semuenzaniso, magetsi, mafuta, uye makemikari) (Pant et al., 2012; Nazari et al., 2020). Kuuya kwe high-throughput genome sequencing uye omics tools/methods kwakapa ruzivo rutsva pamusoro pe genetic regulation, proteomics, uye fluxomics yemaitiro e bacteria dzakasiyana-siyana dzinoora. Kubatanidza maturusi aya ne system biology kwakawedzera kunzwisisa kwedu kusarudzwa uye kugadziriswa kwenzira dze catabolic dzinotarirwa mu bacteria (kureva, metabolic design) kuti uwane biodegradation inoshanda uye inoshanda. Kuti tigadzire nzira dzinoshanda dze bioremediation tichishandisa bacteria dzakakodzera, tinofanira kunzwisisa kugona kwe biochemical, kusiyana kwe metabolic, genetic composition, uye ecology (autoecology/synecology) ye bacteria.
Mufananidzo 1. Manyuko nenzira dzemaPAH asina mamorekuru mashoma kuburikidza nenzvimbo dzakasiyana-siyana dzezvakatipoteredza nezvinhu zvakasiyana-siyana zvinokanganisa biota. Mitsetse ine madhayamita inomiririra kudyidzana pakati pezvinhu zve ecosystem.
Muongororo iyi, takaedza kupfupisa data pamusoro pekuora kwemaPAH akareruka akadai senaphthalene uye naphthalenes dzakatsiviwa nemarudzi akasiyana-siyana emabhakitiriya anofukidza nzira dzemetabolism nekusiyana-siyana, maenzymes anobatanidzwa mukuora, kuumbwa kwemajini/zviri mukati nekusiyana-siyana, mhinduro dzemasero uye zvinhu zvakasiyana-siyana zve bioremediation. Kunzwisisa huwandu hwebiochemical nemamorekuru kuchabatsira mukuona mhando dzakakodzera dzemajini uye kufambira mberi kwavo kwemajini ekugadzirisa zvinosvibisa zvakadaro. Izvi zvichabatsira mukugadzira nzira dzekugadzira nzvimbo chaiyo yemabhakitiriya ekugadzirisa zvinobudirira.
Kuvapo kwehuwandu hwakawanda hwemakemikari ane chepfu uye ane njodzi (zvichigutsa mutemo weHuckel 4n + 2π maerekitironi, n = 1, 2, 3, …) kunotyisidzira zvakanyanya kune zvakasiyana-siyana zvezvakatipoteredza zvakaita semhepo, ivhu, sediments, uye pamusoro pemvura yepasi pevhu (Puglisi et al., 2007). Makemikoro aya ane rings dzebenzene imwe chete (monocyclic) kana dzakawanda benzene rings (polycyclic) dzakarongwa muchimiro chemutsetse, angular kana cluster uye dzinoratidza kugadzikana (kugadzikana/kusagadzikana) munzvimbo nekuda kwesimba rakawanda re resonance uye kusagadzikana (kusagadzikana), izvo zvinogona kutsanangurwa nekusagadzikana kwavo mumvura uye mamiriro akaderera. Kana mhete inonhuhwirira ikatsiviwazve ne methyl (-CH3), carboxyl (-COOH), hydroxyl (-OH), kana sulfonate (-HSO3) mapoka, inova yakasimba, ine hukama hwakasimba hwema macromolecules, uye inounganidzwa muhupenyu hwezvinhu zvipenyu (Seo et al., 2009; Phale et al., 2020). Mamwe ma polycyclic aromatic hydrocarbons (LMWAHs) ane uremu hwakaderera, akadai se naphthalene nezvimwe zvinobva mairi [methylnaphthalene, naphthoic acid, naphthalenesulfonate, uye 1-naphthyl N-methylcarbamate (carbaryl)], akaiswa mune runyorwa rwezvinosvibisa organic neUS Environmental Protection Agency sezvine chepfu, mutagenic, uye/kana carcinogenic (Cerniglia, 1984). Kuburitswa kwerudzi urwu rweNM-PAHs munzvimbo inonaya mvura kunogona kukonzera kuunganidzwa kwemakemikari aya pamatanho ese ekudya, nokudaro zvichikanganisa hutano hwezvinhu zvipenyu (Binkova et al., 2000; Srogi, 2007; Quinn et al., 2009).
Manyuko nenzira dzePAHs kuenda kubiota zvinonyanya kuburikidza nekufamba uye kudyidzana pakati pezvinhu zvakasiyana zve ecosystem zvakaita sevhu, mvura yepasi, mvura yepamusoro, zvirimwa nemhepo (Arey naAtkinson, 2003). Mufananidzo 1 unoratidza kudyidzana uye kugoverwa kwePAHs dzakasiyana dzine huremu hwakaderera mu ecosystem uye nzira dzadzo dzekuenda kubiota/kusangana nevanhu. PAHs dzinoiswa pamusoro nekuda kwekusvibiswa kwemhepo uye kuburikidza nekufamba (kudonha) kwemhepo inoburitsa mota, magasi emuchina wemaindasitiri (kuburitsa gasi remarasha, kupiswa uye kugadzirwa kwecoke) uye kuiswa kwadzo. Mabasa emaindasitiri akadai sekugadzira machira ekugadzira, madhayi nependi; kuchengetedza huni; kugadzira rabha; mabasa ekugadzira simende; kugadzirwa kwezvipembenene; uye kushandiswa kwekurima ndiwo manyuko makuru ePAHs mupasi nepasi pemvura (Bamforth naSingleton, 2005; Wick et al., 2011). Ongororo dzakaratidza kuti ivhu riri munzvimbo dziri pedyo nemaguta, pedyo nemigwagwa mikuru, uye mumaguta makuru rinonyanya kukanganiswa nepolycyclic aromatic hydrocarbons (PAHs) nekuda kweutsi hunobva kumafekitori emagetsi, kupisa kwedzimba, mitoro yemhepo nemigwagwa, uye mabasa ekuvaka (Suman et al., 2016). (2008) yakaratidza kuti maPAH ari muvhu riri pedyo nemigwagwa muNew Orleans, Louisiana, USA aive akakwira kusvika 7189 μg/kg, nepo munzvimbo yakashama, aive 2404 μg/kg chete. Saizvozvowo, huwandu hwePAH hwakakwira kusvika 300 μg/kg hwakataurwa munzvimbo dziri pedyo nenzvimbo dzegesi dzemarasha mumaguta akati wandei eUS (Kanaly naHarayama, 2000; Bamforth naSingleton, 2005). Ivhu rinobva kumaguta akasiyana-siyana eIndia akadai seDelhi (Sharma et al., 2008), Agra (Dubey et al., 2014), Mumbai (Kulkarni naVenkataraman, 2000) uye Visakhapatnam (Kulkarni et al., 2014) rakataurwa kuti rine huwandu hwakawanda hwePAHs. Makemikari anonhuhwirira anonyanya kunamirwa pazvimedu zvevhu, zvinhu zvakasikwa uye zvicherwa zvevhu, nokudaro zvichiva kunyura kukuru kwekabhoni muzvipenyu (Srogi, 2007; Peng et al., 2008). Manyuko makuru ePAHs muzvipenyu zvemvura imvura inonaya (mvura yakanyorova/yakaoma uye utsi hwemvura), mvura inoyerera mumaguta, kuburitswa kwemvura yakasviba, kuburitswa kwemvura yepasi pevhu nezvimwewo. (Srogi, 2007). Zvinofungidzirwa kuti inenge 80% yePAHs muzvipenyu zvegungwa inobva mukunaya kwemvura, kuunganidzwa kwevhu, uye kuburitswa kwetsvina (Motelay-Massei et al., 2006; Srogi, 2007). Kuwanda kwePAH mumvura yepamusoro kana leachate kubva kunzvimbo dzekurasira marara akaoma kunozopedzisira kwanyura mumvura yepasi, zvichikonzera njodzi huru kuhutano hwevanhu sezvo vanhu vanopfuura 70% muSouth neSoutheast Asia vanonwa mvura yepasi (Duttagupta et al., 2019). Ongororo yakaitwa naDuttagupta et al. (2020) yerwizi (32) nemvura yepasi (235) kubva kuWest Bengal, India, yakawana kuti vanhu vanosvika 53% vevagari vemumaguta ne44% yevagari vekumaruwa (vanosvika mamiriyoni 20) vanogona kunge vaine naphthalene (4.9–10.6 μg/L) nezvimwe zvinobva mairi. Mashandisirwo akasiyana evhu uye kuwedzerwa kwekutorwa kwemvura pasi pevhu zvinoonekwa sezvinhu zvikuru zvinodzora kutakurwa kwePAH dzine huremu hwakaderera pasi pevhu. Mvura inoyerera muminda, mvura inobuda mumadhorobha nemaindasitiri, uye tsvina/marara zvakawanikwa zvichikanganiswa nePAH mumigero yerwizi nemadhaka pasi pevhu. Mvura inonaya mumhepo inowedzera kuwedzera kusvibiswa kwePAH. Kuwanda kwePAHs ne alkyl derivatives dzadzo (51 pamwe chete) kwakaonekwa munzizi/nzvimbo dzinoyerera mvura pasi rose, dzakadai seFraser River, Louan River, Denso River, Missouri River, Anacostia River, Ebro River, neDelaware River (Yunker et al., 2002; Motelay-Massei et al., 2006; Li et al., 2010; Amoako et al., 2011; Kim et al., 2018). Mumvura yegungwa reRwizi rweGanges, naphthalene nephenanthrene zvakaonekwa kuti ndizvo zvinonyanya kukosha (zvakawanikwa mu70% yesampuli) (Duttagupta et al., 2019). Uyezve, zvidzidzo zvakaratidza kuti chlorine yemvura yekunwa inogona kutungamira mukuumbwa kwePAHs dzine chepfu uye dzakaiswa chlorine (Manoli naSamara, 1999). MaPAH anoungana muzviyo, michero nemiriwo nekuda kwekutorwa nemiti kubva muvhu rakasviba, pasi pevhu uye mvura inonaya (Fismes et al., 2002). Zvipenyu zvakawanda zvemumvura zvakaita sehove, mussels, clams uye shrimp zvinosvibiswa nemaPAH kuburikidza nekudya chikafu chakasviba nemvura yegungwa, pamwe nenyama neganda (Mackay naFraser, 2000). Nzira dzekubika/kugadzirisa dzakadai sekubika, kugocha, kusvuta, kugocha, kuomesa, kubika uye kubika marasha zvinogonawo kukonzera huwandu hwakawanda hweMAPAH muchikafu. Izvi zvinonyanya kutsamira pasarudzo yezvinhu zvinosvuta, huwandu hwephenolic/aromatic hydrocarbon, maitiro ekubika, rudzi rweheater, huwandu hwehunyoro, oxygen supply uye kupisa kwekupisa (Guillén et al., 2000; Gomes et al., 2013). Polycyclic aromatic hydrocarbons (PAHs) dzakaonekwawo mumukaka pahuwandu hwakasiyana (0.75–2.1 mg/L) (Girelli et al., 2014). Kuunganidzwa kwePAH idzi muchikafu kunoenderanawo nehunhu hwefizikisi yezvekudya, nepo mhedzisiro yadzo ine chepfu ine chekuita nemabasa emuviri, mashandiro emetabolism, kunyudzwa, kugoverwa uye kugoverwa kwemuviri (Mechini et al., 2011).
Chepfu uye mhedzisiro inokuvadza yepolycyclic aromatic hydrocarbons (PAHs) zvave zvichizivikanwa kwenguva yakareba (Cherniglia, 1984). Mapolycyclic aromatic hydrocarbons ane huremu hwakaderera (LMW-PAHs) (marin'i maviri kusvika matatu) anogona kubatana nema macromolecules akasiyana-siyana akadai seDNA, RNA nemapuroteni uye anokonzera carcinogenic (Santarelli et al., 2008). Nekuda kwehunhu hwavo hwekusafarira mvura, anoparadzaniswa ne lipid membranes. Muvanhu, cytochrome P450 monooxygenases inosanganisa maPAHs kuita ma epoxides, mamwe acho anonyanya kuita reactive (semuenzaniso, baediol epoxide) uye anogona kutungamira mukushandurwa kwemasero akajairwa kuita ekenza (Marston et al., 2001). Pamusoro pezvo, zvigadzirwa zvekuchinja zvemaPAH zvakaita sequinones, phenols, epoxides, diols, nezvimwewo zvine chepfu kupfuura zvinoumba masero. Mamwe maPAH nemaitiro awo emetabolism anogona kukanganisa mahormone nema enzyme akasiyana-siyana mumetabolism, nokudaro achikanganisa kukura, central nervous system, masystem ekubereka uye ekudzivirira muviri (Swetha naPhale, 2005; Vamsee-Krishna et al., 2006; Oostingh et al., 2008). Kusangana kwenguva pfupi nePAH dzine huremu hushoma hwemamorekuru kwakataurwa kuti kunokonzera kusashanda zvakanaka kwemapapu uye thrombosis muvanhu vane asthma uye kuwedzera njodzi yekenza yeganda, yemapapu, yedundira uye yemudumbu (Olsson et al., 2010; Diggs et al., 2011). Zvidzidzo zvemhuka zvakaratidzawo kuti kusangana nePAH kunogona kukanganisa mashandiro ekubereka uye kukura uye kunogona kukonzera cataract, kukuvara kweitsvo nechiropa, uye jaundice. Zvigadzirwa zvakasiyana-siyana zvePAH biotransformation zvakaita se diols, epoxides, quinones uye free radicals (cations) zvakaratidzwa kuti zvinoumba DNA adducts. Madhiragi akagadzikana akaratidzwa kuti anochinja michina yekudzokorodza DNA, nepo madhiragi asina kugadzikana anogona kubvisa hurwa hweDNA (kunyanya kuita adenine uye dzimwe nguva kuita guanine); ese ari maviri anogona kugadzira zvikanganiso zvinotungamira kukuchinja (Schweigert et al. 2001). Pamusoro pezvo, maquinones (benzo-/pan-) anogona kugadzira reactive oxygen species (ROS), zvichikonzera kukuvara kunouraya kuDNA nemamwe macromolecules, nokudaro zvichikanganisa mashandiro/kurarama kwetishu (Ewa naDanuta 2017). Kugara uchisangana nehuwandu hwakaderera hwepyrene, biphenyl nenaphthalene kwakataurwa kuti kunokonzera kenza mumhuka dzinoyedzwa (Diggs et al. 2012). Nekuda kwechepfu yavo inouraya, kucheneswa/kubviswa kweaya maPAH kubva panzvimbo dzakakanganiswa/dzakasvibiswa ndicho chinhu chakakosha.
Nzira dzakasiyana-siyana dzemuviri nedzemakemikari dzakashandiswa kubvisa maPAH munzvimbo/nzvimbo dzakasvibiswa. Maitiro akadai sekupiswa, kubviswa kwechlorine, kuoxidation yeUV, kugadzirisa, uye kubvisa solvent ane zvakawanda zvakaipa, zvinosanganisira kugadzirwa kwezvinhu zvine chepfu, kuoma kwemaitiro, matambudziko ekuchengetedza nekutonga, kushanda zvishoma, uye mutengo wakakwira. Zvisinei, kuora kwebhakitiriya (kunonzi bioremediation) inzira inovimbisa inosanganisira kushandiswa kwetupukanana muchimiro chetsika dzakachena kana makoroni. Kana tichienzanisa nenzira dzemuviri nedzemakemikari, maitiro aya haana njodzi kune nharaunda, haapindi, haadhuri, uye anochengetedza nharaunda. Kugadziriswa kwebhakitiriya kunogona kuitwa panzvimbo yakakanganiswa (in situ) kana panzvimbo yakagadzirirwa (ex situ) uye nekudaro kunoonekwa senzira yekugadzirisa yakasimba kupfuura nzira dzechinyakare dzemuviri nedzemakemikari (Juhasz naNaidu, 2000; Andreoni naGianfreda, 2007; Megharaj et al., 2011; Phale et al., 2020; Sarkar et al., 2020).
Kunzwisisa matanho ekuchinja kwemakemikari anokonzera kusvibiswa kwezvinhu zvinonhuhwirira kune zvakunoreva zvikuru mune zvesainzi nehupfumi pakuchengetedza nharaunda nezvakatipoteredza. Kabhoni inofungidzirwa kuti inosvika 2.1 × 1018 magiramu (C) inochengetwa mumadhaka uye makemikari eorganic (kureva, mafuta, gasi rechisikigo, uye marasha, kureva mafuta efossil) pasi rose, zvichiita mupiro mukuru mukutenderera kwekabhoni pasi rose. Zvisinei, kukura nekukurumidza kwemaindasitiri, kubviswa kwemafuta efossil, uye mabasa evanhu zviri kuderedza nzvimbo idzi dzekabhoni dzinochengeterwa lithospheric, zvichiburitsa kabhoni inofungidzirwa kuti inosvika 5.5 × 1015 g (sezvinosvibisa) mumhepo gore rega rega (Gonzalez-Gaya et al., 2019). Kabhoni iyi yakawanda inopinda munzvimbo dzepasi nedzegungwa kuburikidza nekudonha, kutakurwa, uye kuyerera kwemvura. Pamusoro pezvo, tsvina itsva dzinobva mumafuta efossil, dzakadai semapurasitiki, mapurasitiki uye mapurasitiki anosimbisa (phthalates nemaisomers avo), dzinosvibisa zvakanyanya nharaunda dzegungwa, ivhu nedzemumvura uye biota yadzo, nokudaro zvichiwedzera njodzi dzemamiriro ekunze epasi rose. Mhando dzakasiyana dzema microplastics, nanoplastics, zvidimbu zvepurasitiki uye zvigadzirwa zvavo zvine chepfu zvinobva ku polyethylene terephthalate (PET) zvakaungana muPacific Ocean pakati peNorth America neSoutheast Asia, zvichiumba "Great Pacific Garbage Patch", zvichikuvadza zvipenyu zvemugungwa (Newell et al., 2020). Zvidzidzo zvesainzi zvakaratidza kuti hazvigoneke kubvisa tsvina/marara akadaro nenzira dzepanyama kana dzemakemikari. Munyaya iyi, ma microorganisms anobatsira zvikuru ndeaya anokwanisa kushandura tsvina kuita carbon dioxide, chemical energy nezvimwe zvinhu zvisina chepfu zvinozopedzisira zvapinda mune mamwe maitiro ekudyara (H, O, N, S, P, Fe, nezvimwewo). Saka, kunzwisisa hutachiona hwe microorganism hwekusvibisa mineralization uye kudzora kwayo zvakatipoteredza kwakakosha pakuongorora kutenderera kwe microorganism carbon, bhajeti ye net carbon uye njodzi dzemamiriro ekunze mune ramangwana. Zvichienderana nekudiwa kukuru kwekubvisa mishonga yakadaro kubva munharaunda, maindasitiri akasiyana-siyana e eco-industry akatarisana ne clean technologies akabuda. Neimwe nzira, kukoshesa marara emaindasitiri/marara akaunganidzwa muzvipenyu (kureva nzira yekuwara kusvika kupfuma) kunoonekwa seimwe yenheyo dzehupfumi hwakatenderera uye zvinangwa zvebudiriro inoenderera mberi (Close et al., 2012). Nokudaro, kunzwisisa metabolic, enzymes uye majini ezvinogona kukonzera kuora kwezvinhu izvi kwakakosha zvikuru pakubvisa zvinobudirira uye kugadzirisa tsvina yakadaro inonhuhwirira.
Pakati pezvinhu zvakawanda zvinosvibisa zvinhu zvinonhuhwirira, tinonyanya kutarisisa maPAH ane huremu hushoma se naphthalene uye naphthalene dzakatsiviwa. Makemikoro aya ndiwo makuru emafuta anobva mupetroleum, madhayi emachira, zvigadzirwa zvevatengi, mishonga yezvipembenene (mothballs uye zvinodzinga zvipembenene), plasticizers uye tannins uye nekudaro akapararira munzvimbo dzakawanda dzezvakatipoteredza (Preuss et al., 2003). Mishumo ichangoburwa inoratidza kuunganidzwa kwehuwandu hwenaphthalene mumadhaka emvura, pasi pevhu nevhu riri pasi pevhu, nzvimbo dzevadose nemahombekombe erwizi, zvichiratidza kuti inoungana munzvimbo (Duttagupta et al., 2019, 2020). Tafura 2 inopfupikisa hunhu hwefizikisi, mashandisirwo ayo uye mhedzisiro yehutano yenaphthalene nezvimwe zvinobva mairi. Kana tichienzanisa nedzimwe PAH dzine huremu hwakawanda, naphthalene nezvimwe zvinobva mairi hazvina kunyanya kunyungudika mumvura, hazvinyungudike mumvura uye zvinopararira zvakanyanya munzvimbo dzezvakatipoteredza, saka zvinowanzo shandiswa senzira dzekudzidza metabolism, majini uye kusiyana kwemetabolism yePAHs. Huwandu hwakawanda hwetupukanana tunokwanisa kushandura naphthalene nezvinhu zvinobva mairi, uye ruzivo rwakakwana rwunowanikwa nezvenzira dzayo dzekugadzirisa metabolic, ma enzymes uye zvinhu zvinodzora (Mallick et al., 2011; Phale et al., 2019, 2020). Pamusoro pezvo, naphthalene nezvinhu zvinobva mairi zvakasarudzwa sema prototype compounds ekuongorora kusvibiswa kwezvakatipoteredza nekuda kwekuwanda kwazvo uye kuwanikwa kwadzo muhupenyu. US Environmental Protection Agency inofungidzira kuti avhareji yehuwandu hwe naphthalene i5.19 μg pa cubic meter kubva muutsi hwefodya, kunyanya kubva mukutsva kusina kukwana, uye 7.8 kusvika 46 μg kubva muutsi hweparutivi, nepo kusangana ne creosote ne naphthalene kuri pamusoro pe100 kusvika 10,000 (Preuss et al. 2003). Naphthalene kunyanya yakawanikwa ine chepfu yekufema uye kenza yemarudzi, dunhu, uye bonde. Zvichibva pazvidzidzo zvemhuka, International Agency for Research on Cancer (IARC) yakaisa naphthalene mu "inogona kukonzera kenza kuvanhu" (Boka 2B)1. Kusangana ne naphthalenes dzakatsiviwa, kunyanya nekufemerwa kana kupiwa nemuromo, kunokonzera kukuvara kwemapfupa emapapu uye kunowedzera kuwanda kwemabundu emapapu mumakonzo nemakonzo (National Toxicology Program 2). Migumisiro yakaipa inosanganisira kuda kurutsa, kurutsa, kurwadziwa mudumbu, manyoka, musoro, kuvhiringidzika, kudikitira zvakanyanya, fivha, tachycardia, nezvimwewo. Kune rumwe rutivi, mushonga wezvipembenene we carbamate carbaryl (1-naphthyl N-methylcarbamate) wakataurwa kuti une chepfu kune mhuka dzisina mapfupa emumvura, mhuka dzinogara mumvura, nyuchi dzeuchi nevanhu uye zvakaratidzwa kuti zvinodzivisa acetylcholinesterase zvichikonzera kuoma mitezo (Smulders et al., 2003; Bulen naDistel, 2011). Nokudaro, kunzwisisa mashandiro ekuora kwehutachiona, kutonga kwemajini, maitiro enzymatic uye masero kwakakosha pakugadzira nzira dzekugadzirisa bioremediation munzvimbo dzakasvibiswa.
Tafura 2. Ruzivo rwakadzama nezvehunhu hwefizikisi, mashandisirwo, nzira dzekuzivikanwa nadzo uye zvirwere zvine chekuita nenaphthalene nezvinobva mairi.
Munzvimbo dzakasvibiswa, tsvina inonhuhwirira inonyungudutsa mvura uye inonyungudutsa mafuta inogona kukonzera mhedzisiro dzakasiyana-siyana dzemasero pa microbiome yenharaunda (nharaunda), dzakadai sekuchinja kwekuyerera kwemembrane, kubvumidzwa kwemembrane, kuzvimba kwe lipid bilayer, kukanganisa kufambiswa kwesimba (electron transport chain/proton motivation force), uye mashandiro emapuroteni akabatana nemembrane (Sikkema et al., 1995). Pamusoro pezvo, zvimwe zvinonyunguduka senge catechols nequinones zvinogadzira reactive oxygen species (ROS) uye zvinoumba adducts neDNA nemapuroteni (Penning et al., 1999). Saka, kuwanda kwemakomponi akadaro mu ecosystems kunoita kuti nharaunda dzema microbes dzive dzinokanganisa mashandiro emuviri pamazinga akasiyana-siyana emuviri, kusanganisira kutorwa/kutakurwa, kushandurwa kwemukati memasero, kushandiswa/kushandiswa, uye kupatsanurwa.
Kutsvaga kweRibosomal Database Project-II (RDP-II) kwakaratidza kuti mhando dzebhakitiriya dzinosvika 926 dzakaparadzaniswa kubva mumarudzi ezvirimwa kana kuti enhancement cultures dzakasvibiswa ne naphthalene kana zvinobva mairi. Boka reProteobacteria ndiro raiva nehuwandu hwakanyanya hwevamiririri (n = 755), richiteverwa neFirmicutes (52), Bacteroidetes (43), Actinobacteria (39), Tenericutes (10), uye mabhakitiriya asina kurongwa (8) (Mufananidzo 2). Vamiririri veγ-Proteobacteria (Pseudomonadales neXanthomonadales) vakatonga mapoka ese eGram-negative ane huwandu hwakawanda hweG+C (54%), nepo Clostridiales neBacillales (30%) aive mapoka eGram-positive ane huwandu hwakaderera hweG+C. Pseudomonas (nhamba yepamusoro, mhando 338) dzakanzi dzinokwanisa kuparadza naphthalene nezvinhu zvinobva mairi mune zvakasiyana-siyana zvakasvibiswa (tara yemarasha, peturu, mafuta asina kuchena, tsvina, mafuta akadururwa, mvura yakasviba, marara eorganic nenzvimbo dzakarasirwa marara) pamwe chete nenzvimbo dzisina kusvibiswa (ivhu, nzizi, madhaka nemvura yepasi pevhu) (Mufananidzo 2). Uyezve, zvidzidzo zvekuwedzera uye ongororo yemetagenomic yemamwe matunhu aya zvakaratidza kuti mhando dzeLegionella neClostridium dzisina kurimwa dzinogona kuva nekugona kuora, zvichiratidza kudiwa kwekurima mabhakitiriya aya kuti adzidze nzira itsva uye kusiyana kwemetabolism.
Mufananidzo 2. Kusiyana-siyana kwetaxonomic uye kupararira kwebhakitiriya munzvimbo dzakasvibiswa ne naphthalene uye naphthalene derivatives.
Pakati pezvipembenene zvakasiyana-siyana zvinoparadza hydrocarbon, zvakawanda zvinokwanisa kukanganisa naphthalene senzvimbo yega inopa kabhoni nesimba. Kurongeka kwezviitiko zvine chekuita ne metabolism ye naphthalene kwakatsanangurwa nezvePseudomonas sp. (marudzi: NCIB 9816-4, G7, AK-5, PMD-1 naCSV86), Pseudomonas stutzeri AN10, Pseudomonas fluorescens PC20 nedzimwe mhando (ND6 naAS1) (Mahajan et al., 1994; Resnick et al., 1996; Annweiler et al., 2000; Basu et al., 2003; Dennis naZylstra, 2004; Sota et al., 2006; Metabolism inotangwa ne multicomponent dioxygenase [naphthalene dioxygenase (NDO), ring hydroxylating dioxygenase] iyo inokonzeresa oxidation yeimwe ye aromatic rings ye naphthalene ichishandisa molecular oxygen seimwe substrate, ichishandura naphthalene kuita cis-naphthalenediol (Mufananidzo 3). Cis-dihydrodiol inoshandurwa kuita 1,2-dihydroxynaphthalene ne dehydrogenase. Dioxygenase inocheka ring, 1,2-dihydroxynaphthalene dioxygenase (12DHNDO), inoshandura 1,2-dihydroxynaphthalene kuita 2-hydroxychromene-2-carboxylic acid. Enzymatic cis-trans isomerization inogadzira trans-o-hydroxybenzylidenepyruvate, iyo inobviswa nehydratase aldolase kuita salicylic aldehyde nepyruvate. Organic acid pyruvate ndiyo yaive yekutanga C3 compound yakabva mu naphthalene carbon skeleton ndokuendeswa munzira yepakati pekabhoni. Pamusoro pezvo, NAD+-dependent salicylaldehyde dehydrogenase inoshandura salicylaldehyde kuita salicylic acid. Metabolism padanho iri inonzi "nzira yepamusoro" yekuora kwe naphthalene. Nzira iyi yakajairika mumabhakitiriya mazhinji anoora ne naphthalene. Zvisinei, kune mashoma mashoma; semuenzaniso, mu thermophilic Bacillus hamburgii 2, kuora kwe naphthalene kunotanga ne naphthalene. 2,3-dioxygenase kuumba 2,3-dihydroxynaphthalene (Annweiler et al., 2000).
Mufananidzo 3. Nzira dze naphthalene, methylnaphthalene, naphthoic acid, uye carbaryl degradation. Nhamba dzakatenderedzwa dzinomiririra ma enzymes anokonzera shanduko yakatevedzana ye naphthalene nezvimwe zvinobva mairi kuita zvigadzirwa zvinotevera. 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.
Zvichienderana nehupenyu uye majini ahwo, salicylic acid inobuda inowedzerwa kushandiswa kuburikidza ne catechol pathway uchishandisa salicylate 1-hydroxylase (S1H) kana kuburikidza ne gentisate pathway uchishandisa salicylate 5-hydroxylase (S5H) (Mufananidzo 3). Sezvo salicylic acid iri iyo inonyanya kushandiswa mu naphthalene metabolism (nzira yepamusoro), matanho kubva ku salicylic acid kuenda kuTCA intermediate anowanzo kunzi nzira yepasi, uye majini akarongwa kuita operon imwe chete. Zvakajairika kuona kuti majini ari mu upper pathway operon (nah) uye lower pathway operon (sal) anodzorwa nezvinhu zvinodzorwa; semuenzaniso, NahR uye salicylic acid zvinoshanda se inducers, zvichibvumira ma operon ese kuti ashandise naphthalene zvizere (Phale et al., 2019, 2020).
Pamusoro pezvo, catechol inonamirwa cyclically ku 2-hydroxymuconate semialdehyde kuburikidza nenzira ye meta ne catechol 2,3-dioxygenase (C23DO) (Yen et al., 1988) uye inowedzerwa hydrolyzed ne 2-hydroxymuconate semialdehyde hydrolase kuti igadzire 2-hydroxypent-2,4-dienoic acid. 2-hydroxypent-2,4-dienoate inozoshandurwa kuita pyruvate neacetaldehyde ne hydratase (2-oxopent-4-enoate hydratase) uye aldolase (4-hydroxy-2-oxopentanoate aldolase) uye yozopinda munzira yepakati pekabhoni (Mufananidzo 3). Neimwe nzira, catechol inonamirwa cyclically ku cis,cis-muconate kuburikidza nenzira ye ortho ne catechol 1,2-oxygenase (C12DO). Muconate cycloisomerase, muconolactone isomerase, uye β-ketoadipate-nollactone hydrolase zvinoshandura cis,cis-muconate kuita 3-oxoadipate, iyo inopinda munzira yepakati pekabhoni kuburikidza nesuccinyl-CoA uye acetyl-CoA (Nozaki et al., 1968) (Mufananidzo 3).
Munzira ye gentisate (2,5-dihydroxybenzoate), mhete inonhuhwirira inotsemurwa ne gentisate 1,2-dioxygenase (GDO) kuti igadzire maleylpyruvate. Chigadzirwa ichi chinogona kusanganiswa zvakananga kuti chiite pyruvate uye malate, kana kuti chinogona kusanganiswa kuti chiite fumarylpyruvate, iyo inogona kusanganiswa kuti iite pyruvate uye fumarate (Larkin and Day, 1986). Sarudzo yenzira imwe yakaonekwa mumabhakitiriya eGram-negative neGram-positive pamazinga ebiochemical ne genetic (Morawski et al., 1997; Whyte et al., 1997). Mabhakitiriya eGram-negative (Pseudomonas) anosarudza kushandisa salicylic acid, iyo inokonzera metabolism ye naphthalene, ichiibvisa mucarboxylating kuita catechol vachishandisa salicylate 1-hydroxylase (Gibson naSubramanian, 1984). Kune rumwe rutivi, mubhakitiriya reGram-positive (Rhodococcus), salicylate 5-hydroxylase inoshandura salicylic acid kuita gentisic acid, nepo salicylic acid isina inductive effect pakunyorwa kwemajini e naphthalene (Grund et al., 1992) (Mufananidzo 3).
Zvakataurwa kuti mhando dzakadai sePseudomonas CSV86, Oceanobacterium NCE312, Marinhomonas naphthotrophicus, Sphingomonas paucimobilis 2322, Vibrio cyclotrophus, Pseudomonas fluorescens LP6a, Pseudomonas uye Mycobacterium species dzinogona kukanganisa monomethylnaphthalene kana dimethylnaphthalene (Dean-Raymond naBartha, 1975; Cane naWilliams, 1982; Mahajan et al., 1994; Dutta et al., 1998; Hedlund et al., 1999). Pakati pavo, nzira ye1-methylnaphthalene uye 2-methylnaphthalene degradation pathway yePseudomonas sp. CSV86 yakadzidzwa zvakajeka pamazinga ebiochemical uye enzyme (Mahajan et al., 1994). 1-Methylnaphthalene inogadziriswa kuburikidza nenzira mbiri. Kutanga, mhete inonhuhwirira inogadzirwa nehydroxylated (mhete isina kutsiviwa yemethylnaphthalene) kuti igadzire cis-1,2-dihydroxy-1,2-dihydro-8-methylnaphthalene, iyo inozowedzerwa kuita methyl salicylate uye methylcatechol, uye yozopinda munzira yepakati yekabhoni mushure mekutsemuka kweringi (Mufananidzo 3). Nzira iyi inonzi "nzira yekabhoni". Munzira yechipiri ye "detoxification", boka remethyl rinogona kuwedzeredzwa neNDO kuti rigadzire 1-hydroxymethylnaphthalene, iyo inowedzeredzwa kuita 1-naphthoic acid uye inoburitswa munzvimbo yekusimwa sechinhu chisina simba. Zvidzidzo zvakaratidza kuti strain CSV86 haigone kukura pa 1- uye 2-naphthoic acid senzvimbo yega yekabhoni nesimba, zvichisimbisa nzira yayo yekubvisa detoxification (Mahajan et al., 1994; Basu et al., 2003). Mu 2-methylnaphthalene, boka re methyl rinoitwa hydroxylation ne hydroxylase kuti rigadzire 2-hydroxymethylnaphthalene. Pamusoro pezvo, rin'i isina kutsiviwa yerin'i re naphthalene inoitwa rin'i re hydroxylation kuti igadzire dihydrodiol, iyo inogadzirwa kuita 4-hydroxymethylcatechol muzvikamu zvakasiyana-siyana zve enzyme-catalyzed reactions uye inopinda munzira yepakati pekabhoni kuburikidza nenzira ye meta-ring cleavage. Saizvozvowo, S. paucimobilis 2322 yakataurwa kuti inoshandisa NDO kuita hydroxylate 2-methylnaphthalene, iyo inogadzirwazve kuita methyl salicylate uye methylcatechol (Dutta et al., 1998).
Naphthoic acids (yakatsiviwa/isina kutsiviwa) imishonga inobvisa chepfu mumuviri/biotransformation inogadzirwa panguva yekuora kwemethylnaphthalene, phenanthrene neanthracene uye inoburitswa munzvimbo inocherwa zvirimwa. Zvakataurwa kuti Stenotrophomonas maltophilia CSV89 inokwanisa kugaya 1-naphthoic acid senzvimbo yekabhoni (Phale et al., 1995). Metabolism inotanga ne dihydroxylation ye aromatic ring kuti igadzire 1,2-dihydroxy-8-carboxynaphthalene. Diol inobuda inosanganiswa kuita catechol kuburikidza ne 2-hydroxy-3-carboxybenzylidenepyruvate, 3-formylsalicylic acid, 2-hydroxyisophthalic acid uye salicylic acid uye inopinda munzira yepakati yekabhoni kuburikidza nenzira ye meta-ring cleavage (Mufananidzo 3).
Carbaryl mushonga wezvipembenene une naphthyl carbamate. Kubva paGreen Revolution muIndia muma1970, kushandiswa kwemishonga yemakemikari nemishonga yezvipembenene kwakaita kuti pave nekuwanda kwemafuta anonhuhwirira (PAH) kubva muzvirimwa zvisina nzvimbo (Pingali, 2012; Duttagupta et al., 2020). Inofungidzirwa kuti 55% (85,722,000 hectares) yeminda yese yekurima muIndia inorapwa nemishonga yezvipembenene ine mishonga yekuuraya zvipembenene. Mumakore mashanu apfuura (2015–2020), chikamu chekurima muIndia chakashandisa avhareji yematani 55,000 kusvika 60,000 emishonga yekuuraya zvipembenene pagore (Dhipatimendi reMakambani neKugara zvakanaka kweVarimi, Ministry of Agriculture, Government of India, Nyamavhuvhu 2020). Munzvimbo dzekuchamhembe nepakati peGangetic (nyika dzine huwandu hwevanhu vakawanda uye huwandu hwevanhu), kushandiswa kwemishonga yekuuraya zvipembenene pazvirimwa kwakapararira, mishonga yekuuraya zvipembenene ichinyanya kupararira. Carbaryl (1-naphthyl-N-methylcarbamate) mushonga wezvipembenene une chepfu yakawanda, une chepfu iri pakati nepakati kusvika kune yakanyanya unoshandiswa mukurima kweIndia paavhareji yematani 100–110. Unowanzo kutengeswa pasi pezita rekutengeserana rekuti Sevin uye unoshandiswa kudzora zvipembenene (maaphid, masvosve emoto, nhata, zvikwekwe, madandemutande nezvimwe zvipembenene zvakawanda zvekunze) zvichikanganisa zvirimwa zvakasiyana-siyana (chibage, soya, donje, michero nemiriwo). Zvimwe zvipembenene zvakaita sePseudomonas (NCIB 12042, 12043, C4, C5, C6, C7, Pseudomonas putida XWY-1), Rhodococcus (NCIB 12038), Sphingobacterium spp. (CF06), Burkholderia (C3), Micrococcus neArthrobacter zvinogonawo kushandiswa kudzora zvimwe zvipembenene. Zvakataurwa kuti RC100 inogona kukanganisa carbaryl (Larkin naDay, 1986; Chapalamadugu naChaudhry, 1991; Hayatsu et al., 1999; Swetha naPhale, 2005; Trivedi et al., 2017). Nzira yekuora kwecarbaryl yakaongororwa zvakanyanya pamazinga ebiochemical, enzyme uye genetic muivhu raPseudomonas sp. Strains C4, C5 naC6 (Swetha naPhale, 2005; Trivedi et al., 2016) (Mufananidzo 3). Nzira yekuchinja kwemuviri inotanga nehydrolysis ye ester bond ne carbaryl hydrolase (CH) kuti igadzire 1-naphthol, methylamine uye carbon dioxide. 1-naphthol inozoshandurwa kuita 1,2-dihydroxynaphthalene ne 1-naphthol hydroxylase (1-NH2), iyo inozowedzerwa kushandiswa kuburikidza nenzira yepakati pekabhoni kuburikidza ne salicylate ne gentisate. Mamwe mabhakitiriya anoparadza carbaryl anonzi anoichinja kuita salicylic acid kuburikidza nekutsemuka kwe catechol ortho ring (Larkin na Day, 1986; Chapalamadugu na Chaudhry, 1991). Zvinonyanya kukosha kuti mabhakitiriya anoparadza naphthalene anonyanya kushandisa salicylic acid kuburikidza ne catechol, nepo mabhakitiriya anoparadza carbaryl anofarira kushandisa salicylic acid kuburikidza nenzira ye gentisate.
Naphthalenesulfonic acid/disulfonic acid uye naphthylaminesulfonic acid derivatives zvinogona kushandiswa sepakati pekugadzira azo dyes, wetting agents, dispersants, nezvimwewo. Kunyangwe mishonga iyi isina chepfu yakawanda kuvanhu, kuongororwa kwe cytotoxicity kwakaratidza kuti inouraya hove, daphnia uye algae (Greim et al., 1994). Vamiriri ve genus Pseudomonas (marudzi A3, C22) vakashumwa kuti vanotanga metabolism ne double hydroxylation ye aromatic ring ine sulfonic acid group kuti igadzire dihydrodiol, iyo inozoshandurwazve kuita 1,2-dihydroxynaphthalene nekupatsanuka kwe sulfite group (Brilon et al., 1981). 1,2-dihydroxynaphthalene inobuda inogadziriswa kuburikidza nenzira yekare ye naphthalene, kureva, nzira ye catechol kana gentisate (Mufananidzo 4). Zvakaratidzwa kuti aminonaphthalenesulfonic acid uye hydroxynaphthalenesulfonic acid zvinogona kukanganiswa zvachose nekubatanidzwa kwemabhakitiriya akasiyana-siyana nenzira dzinoenderana dze catabolic (Nortemann et al., 1986). Zvakaratidzwa kuti nhengo imwe chete ye consortium inobvisa aminonaphthalenesulfonic acid kana hydroxynaphthalenesulfonic acid ne 1,2-dioxygenation, nepo aminosalicylate kana hydroxysalicylate ichiburitswa munzvimbo yekukudziridza se dead-end metabolite uye inozotorwa nedzimwe nhengo dze consortium. Naphthalenedisulfonic acid ine polar asi haigone kuora uye saka inogona kugayiwa nenzira dzakasiyana. Kubviswa kwesulfurization kwekutanga kunoitika panguva ye regioselective dihydroxylation ye aromatic ring ne sulfonic acid group; Kubviswa kwechipiri kwesulfurization kunoitika panguva yehydroxylation ye5-sulfosalicylic acid nesalicylic acid 5-hydroxylase kuti igadzire gentisic acid, iyo inopinda munzira yepakati pekabhoni (Brilon et al., 1981) (Mufananidzo 4). Ma enzymes anokonzera kukanganiswa kwe naphthalene ndiwo anokonzerawo metabolism ye naphthalene sulfonate (Brilon et al., 1981; Keck et al., 2006).
Mufananidzo 4. Nzira dzekuchinja kwemuviri dzinoita kuti naphthalene sulfonate iparare. Nhamba dziri mukati memadenderedzwa dzinomiririra ma enzymes anokonzera metabolism ye naphthyl sulfonate, akafanana/akafanana nema enzymes anotsanangurwa muMufananidzo 3.
MaPAH ane huremu hwakaderera (LMW-PAHs) anogona kuderedzwa, haadi mvura uye haanyungudike zvakanaka, saka haakwanise kuputsika/kuora kwechisikigo. Zvisinei, zvipembenene zve aerobic zvinokwanisa kuioxidation nekutora molecular oxygen (O2). Ma enzyme aya anonyanya kuve eboka re oxidoreductases uye anogona kuita maitiro akasiyana-siyana akadai se aromatic ring hydroxylation (mono- kana dihydroxylation), dehydrogenation uye aromatic ring cleavage. Zvigadzirwa zvinowanikwa kubva mumaitiro aya zviri mu oxidation state yakakwira uye zvinogadziriswa zviri nyore kuburikidza ne central carbon pathway (Phale et al., 2020). Ma enzyme ari munzira yekuora anonzi haagone kunyungudika. Basa rema enzyme aya rakaderera zvikuru kana kuti harina basa kana masero akakurira panzvimbo dziri nyore dzekabhoni dzakadai se glucose kana organic acids. Tafura 3 inopfupikisa ma enzyme akasiyana-siyana (oxygenases, hydrolases, dehydrogenases, oxidases, nezvimwewo) anobatanidzwa mu metabolism ye naphthalene nezvimwe zvinobva mairi.
Tafura 3. Hunhu hwemakemikari emakemikari anokonzera kuparara kwe naphthalene nezvinobva mairi.
Zvidzidzo zveRadioisotope (18O2) zvakaratidza kuti kuiswa kwemolecular O2 mu aromatic rings ne oxygenases ndiyo danho rakakosha mukuita kuti pave nekuora kwechinhu (Hayaishi et al., 1955; Mason et al., 1955). Kubatanidzwa kweatomu imwe chete yeokisijeni (O) kubva ku molecular oxygen (O2) mu substrate kunotangwa ne endogenous kana exogenous monooxygenases (inonziwo hydroxylases). Imwe atomu yeokisijeni inoderedzwa kuita mvura. Exogenous monooxygenases inoderedza flavin neNADH kana NADPH, nepo mu endomonooxygenases flavin inoderedzwa ne substrate. Nzvimbo yehydroxylation inokonzera kusiyana mukuumbwa kwechigadzirwa. Semuenzaniso, salicylate 1-hydroxylase hydroxylates salicylic acid panzvimbo yeC1, ichigadzira catechol. Kune rumwe rutivi, salicylate 5-hydroxylase ine zvikamu zvakawanda (ine reductase, ferredoxin, uye oxygenase subunits) hydroxylates salicylic acid panzvimbo yeC5, zvichigadzira gentisic acid (Yamamoto et al., 1965).
MaDioxygenases anosanganisira maatomu maviri eO2 muchikamu chepasi. Zvichienderana nezvigadzirwa zvakagadzirwa, akakamurwa kuita ma ring hydroxylating dioxygenases uye ma ring cleaving dioxygenases. Ma ring hydroxylating dioxygenases anoshandura ma aromatic substrates kuita cis-dihydrodiols (semuenzaniso, naphthalene) uye akapararira mumabhakitiriya. Kusvika pari zvino, zvakaratidzwa kuti zvipenyu zvine ma ring hydroxylating dioxygenases zvinokwanisa kukura pamhando dzakasiyana dze aromatic carbon sources, uye ma enzymes aya akarongwa seNDO (naphthalene), toluene dioxygenase (TDO, toluene), uye biphenyl dioxygenase (BPDO, biphenyl). Zvose NDO neBPDO zvinogona kukonzera double oxidation uye side chain hydroxylation ye polycyclic aromatic hydrocarbons dzakasiyana-siyana (toluene, nitrotoluene, xylene, ethylbenzene, naphthalene, biphenyl, fluorene, indole, methylnaphthalene, naphthalenesulfonate, phenanthrene, anthracene, acetophenone, nezvimwewo) (Boyd naSheldrake, 1998; Phale et al., 2020). NDO isystem ine zvikamu zvakawanda ine oxidoreductase, ferredoxin, uye chikamu chinoshanda chine oxygenase (Gibson naSubramanian, 1984; Resnick et al., 1996). Chikamu chinobatsira cheNDO chine α subunit hombe uye β subunit diki yakarongwa mu α3β3 configuration. NDO ndeyemhuri yakakura yema oxygenases uye α-subunit yayo ine Rieske site [2Fe-2S] uye mononuclear non-heme iron, iyo inosarudza substrate specificity yeNDO (Parales et al., 1998). Kazhinji, mune imwe catalytic cycle, maerekitironi maviri kubva mukuderedza pyridine nucleotide anotamisirwa kuFe(II) ion munzvimbo inoshanda kuburikidza ne reductase, ferredoxin uye Rieske site. Kuderedza equivalents kunoshanda molecular oxygen, inova chinhu chinodiwa kuti substrate dihydroxylation (Ferraro et al., 2005). Kusvika pari zvino, maNDO mashoma chete akacheneswa uye akatsanangurwa zvakadzama kubva kumarudzi akasiyana uye kutonga kwemajini kwenzira dzinobatanidzwa mukuora kwe naphthalene kwakadzidzwa zvakadzama (Resnick et al., 1996; Parales et al., 1998; Karlsson et al., 2003). Ma "ring-cleaving dioxygenases" (endo- kana ortho-ring-cleaving enzymes uye exodiol- kana meta-ring-cleaving enzymes) anoshanda pamakemikari aromatic aromatic anokonzerwa nehydroxylated. Semuenzaniso, "ortho-ring-cleaving dioxygenase" inonzi catechol-1,2-dioxygenase, nepo "meta-ring-cleaving dioxygenase" iri catechol-2,3-dioxygenase (Kojima et al., 1961; Nozaki et al., 1968). Kuwedzera kune akasiyana-siyana ma "oxygenases", kunewo akasiyana-siyana ma "dehydrogenases" anokonzera "dehydrogenation" ye "aromatic dihydrodiols", "alcohols" uye "aldehydes" uye kushandisa NAD+/NADP+ se "electron acceptors", ayo ari mamwe ema "enzymes" akakosha anobatanidzwa mu "metabolism" (Gibson naSubramanian, 1984; Shaw naHarayama, 1990; Fahle et al., 2020).
Ma enzyme akadai se hydrolases (esterases, amidases) iboka rechipiri rakakosha rema enzymes anoshandisa mvura kupatsanura ma covalent bonds uye kuratidza broad substrate specificity. Carbaryl hydrolase nedzimwe hydrolases dzinoonekwa sezvikamu zve periplasm (transmembrane) munhengo dzeGram-negative bacteria (Kamini et al., 2018). Carbaryl ine amide ne ester linkage; saka, inogona ku hydrolyzed ne esterase kana amidase kuti igadzire 1-naphthol. Carbaryl mu Rhizobium rhizobium strain AC10023 uye Arthrobacter strain RC100 zvakanzi zvinoshanda se esterase ne amidase, zvichiteerana. Carbaryl muArthrobacter strain RC100 inoshandawo se amidase. RC100 yakaratidzwa kuti inonyungudutsa mishonga mina yezvipembenene yerudzi rweN-methylcarbamate yakaita secarbaryl, methomyl, mefenamic acid uye XMC (Hayaatsu et al., 2001). Zvakataurwa kuti CH muPseudomonas sp. C5pp inogona kushanda pacarbaryl (100% activity) uye 1-naphthyl acetate (36% activity), asi kwete pa1-naphthylacetamide, zvichiratidza kuti iesterase (Trivesi et al., 2016).
Zvidzidzo zve biochemical, maitiro ekugadzirisa ma enzyme, uye ongororo yemajini zvakaratidza kuti majini ekuora kwe naphthalene ane mayuniti maviri ekutonga anoshandiswa kana kuti "operon": nah (nzira yekukwira, kushandura naphthalene kuita salicylic acid) uye sal (nzira yekudzika, kushandura salicylic acid kuita nzira yepakati yekabhoni kuburikidza ne catechol). Salicylic acid nezvimwe zvakafanana nayo zvinogona kushanda sezvinokonzera (Shamsuzzaman naBarnsley, 1974). Kana paine glucose kana organic acids, operon inodzvinyirirwa. Mufananidzo 5 unoratidza kurongeka kwemajini ese ekuora kwe naphthalene (muchimiro che operon). Mhando dzakasiyana-siyana dzakatumidzwa zita re nah gene (ndo/pah/dox) dzakatsanangurwa uye dzakawanikwa dziine homology yepamusoro (90%) pakati pemarudzi ese ePseudomonas (Abbasian et al., 2016). Majini enzira yekukwira ye naphthalene aiwanzo kurongwa nenzira yekubvumirana sezvakaratidzwa muMufananidzo 5A. Rimwe jini, nahQ, rakataurwawo kuti rine chekuita ne metabolism ye naphthalene uye raiwanzo kuwanikwa pakati pe nahC ne nahE, asi basa raro chairo harisati rajekeswa. Saizvozvowo, jini re nahY, iro rinokonzera chemotaxis inonzwa naphthalene, rakawanikwa kumagumo ekupedzisira e nah operon mune dzimwe nhengo. MuRalstonia sp., jini reU2 rinobatanidza glutathione S-transferase (gsh) rakawanikwa riri pakati pe nahAa ne nahAb asi harina kukanganisa hunhu hwekushandiswa kwe naphthalene (Zylstra et al., 1997).
Mufananidzo 5. Kurongeka kwemajini uye kusiyana-siyana kwakaonekwa panguva yekuora kwenaphthalene pakati pemarudzi emabhakitiriya; (A) Nzira yepamusoro yenaphthalene, kushandiswa kwenaphthalene kuita salicylic acid; (B) Nzira yakaderera yenaphthalene, salicylic acid kuburikidza necatechol kuenda kunzira yepakati yekabhoni; (C) salicylic acid kuburikidza negentisate kuenda kunzira yepakati yekabhoni.
"Nzira yepasi" (sal operon) inowanzo kuve ne nahGTHINLMOKJ uye inoshandura salicylate kuita pyruvate uye acetaldehyde kuburikidza ne catechol metaring cleavage pathway. Jini re nahG (encoding salicylate hydroxylase) rakawanikwa rakachengetwa pamucheto we proximal we operon (Mufananidzo 5B). Zvichienzaniswa nedzimwe mhando dze naphthalene-degrading, mu P. putida CSV86 nah ne sal operon dzakabatana uye dzakabatana zvakanyanya (inenge 7.5 kb). Mune mamwe mabhakitiriya eGram-negative, akadai seRalstonia sp. U2, Polaromonas naphthalenivorans CJ2, uye P. putida AK5, naphthalene inogadzirwa se central carbon metabolite kuburikidza ne gentisate pathway (muchimiro che sgp/nag operon). Kaseti yemajini inowanzo miririrwa muchimiro che nagAaGHabAcAdBFCQEDJI, uko nagR (inokodha LysR-type regulator) iri kumucheto wepamusoro (Mufananidzo 5C).
Carbaryl inopinda muchikamu chepakati chekabhoni kuburikidza nemetabolism ye1-naphthol, 1,2-dihydroxynaphthalene, salicylic acid, uye gentisic acid (Mufananidzo 3). Zvichibva pazvidzidzo zvemajini uye metabolic, zvakakurudzirwa kupatsanura nzira iyi kuita "upstream" (kushandurwa kwecarbaryl kuita salicylic acid), "pakati" (kushandurwa kwesalicylic acid kuita gentisic acid), uye "downstream" (kushandurwa kwegentisic acid kuita central carbon pathway intermediates) (Singh et al., 2013). Kuongororwa kweC5pp (supercontig A, 76.3 kb) kwakaratidza kuti jini remcbACBDEF rinobatanidzwa mukushandurwa kwecarbaryl kuita salicylic acid, zvichiteverwa nemcbIJKL mukushandurwa kwesalicylic acid kuita gentisic acid, uye mcbOQP mukushandurwa kwegentisic acid kuita central carbon intermediates (fumarate nepyruvate, Trivedi et al., 2016) (Mufananidzo 6).
Zvakataurwa kuti ma enzymes anobatanidzwa mukuora kwema hydrocarbon aromatic (kusanganisira naphthalene ne salicylic acid) anogona kukonzerwa nemakomikari anoenderana uye kudziviswa nezvinhu zviri nyore zvekabhoni zvakaita se glucose kana organic acids (Shingler, 2003; Phale et al., 2019, 2020). Pakati penzira dzakasiyana dze metabolism ye naphthalene nezvinobva mairi, maitiro ekutonga e naphthalene ne carbaryl akaongororwa kusvika pamwero wakati. Kune naphthalene, majini ari munzira dzekumusoro nedzepasi anodzorwa ne NahR, LysR-type trans-acting positive regulator. Inodiwa kuti gene re nah riiswe ne salicylic acid uye kuratidzwa kwayo kwepamusoro-soro kunotevera (Yen naGunsalus, 1982). Uyezve, zvidzidzo zvakaratidza kuti integrative host factor (IHF) ne XylR (sigma 54-dependent transcriptional regulator) zvakakoshawo pakuita kuti majini ashande mu naphthalene metabolism (Ramos et al., 1997). Zvidzidzo zvakaratidza kuti ma enzymes e catechol meta-ring opening pathway, kureva catechol 2,3-dioxygenase, anokonzerwa kana paine naphthalene uye/kana salicylic acid (Basu et al., 2006). Zvidzidzo zvakaratidza kuti ma enzymes e catechol ortho-ring opening pathway, kureva catechol 1,2-dioxygenase, anokonzerwa kana paine benzoic acid uye cis,cis-muconate (Parsek et al., 1994; Tover et al., 2001).
MuC5pp, majini mashanu, mcbG, mcbH, mcbN, mcbR uye mcbS, anoronga ma regulators ari emhuri yeLysR/TetR ye transcriptional regulators ane basa rekudzora carbaryl degradation. Gene re homologous mcbG rakawanikwa riine hukama hwakasimba neLysR-type regulator PhnS (58% amino acid identity) inobatanidzwa mu phenanthrene metabolism muBurkholderia RP00725 (Trivesi et al., 2016). Gene re mcbH rakawanikwa riine chekuita ne intermediate pathway (kushandurwa kwe salicylic acid kuita gentisic acid) uye riri reLysR-type transcriptional regulator NagR/DntR/NahR muPseudomonas neBurkholderia. Nhengo dzemhuri iyi dzakanzi dzinoziva salicylic acid se specific effector molecule yekusimudzira majini degradation. Kune rumwe rutivi, majini matatu, mcbN, mcbR uye mcbS, ari eLysR neTetR type transcriptional regulators, akaonekwa munzira yekudzika (genisate-central carbon pathway metabolites).
Mumaprokaryotes, maitiro ekutamisa majini akatenderera (kuwanikwa, kuchinjana, kana kutamiswa) kuburikidza nemaplasmids, transposons, prophages, genomic islands, uye integrative conjugative elements (ICE) ndizvo zvinokonzera plasticity muma genomes ebhakitiriya, zvichikonzera kuwana kana kurasikirwa nemabasa/hunhu hwakati. Izvi zvinobvumira mabhakitiriya kuti akwanise kuchinjika nekukurumidza kune mamiriro akasiyana ezvakatipoteredza, zvichipa mukana wekugadzirisa metabolic kune host, sekuora kwemakemikari aromatic. Shanduko dzemetabolism dzinowanzo kuwanikwa kuburikidza nekugadzirisa maoperon ekuora, nzira dzawo dzekutonga, uye enzyme specificities, izvo zvinoita kuti pave nekuora kwemhando dzakasiyana dzemakemikari aromatic (Nojiri et al., 2004; Phale et al., 2019, 2020). Ma gene cassette ekuora kwe naphthalene akawanikwa ari pamhando dzakasiyana dzezvinhu zvinofamba senge maplasmids (conjugative ne non-conjugative), transposons, genomes, ICEs, uye musanganiswa wemarudzi akasiyana ebhakitiriya (Mufananidzo 5). MuPseudomonas G7, ma nah ne sal operon eplasmid NAH7 anonyorwa nenzira imwe chete uye chikamu che transposon isina kunaka inoda transposase Tn4653 kuti ifambiswe (Sota et al., 2006). MuPseudomonas strain NCIB9816-4, jini iri rakawanikwa pa conjugative plasmid pDTG1 sema operon maviri (anenge 15 kb akaparadzana) ayo akanyorwa nenzira dzakasiyana (Dennis naZylstra, 2004). MuPseudomonas putida strain AK5, non-conjugative plasmid pAK5 inochengetedza enzyme inokonzera naphthalene degradation kuburikidza ne gentisate pathway (Izmalkova et al., 2013). MuPseudomonas strain PMD-1, nah operon iri pachromosome, nepo sal operon iri pa conjugative plasmid pMWD-1 (Zuniga et al., 1981). Zvisinei, muPseudomonas stutzeri AN10, majini ese e naphthalene degradation (nah uye sal operon) ari pachromosome uye anofungidzirwa kuti anotorwa kuburikidza ne transposition, recombination, uye rearrangement events (Bosch et al., 2000). MuPseudomonas sp. CSV86, nah uye sal operon dziri mu genome muchimiro cheICE (ICECSV86). Chimiro ichi chakachengetedzwa ne tRNAGly chinoteverwa nekudzokorora zvakananga kunoratidza nzvimbo dzekubatanidza/kubatanidza (attR ne attL) uye integrase yakafanana ne phage iri kumativi ese e tRNAGly, nokudaro yakafanana ne ICEclc element (ICEclcB13 mu Pseudomonas knackmusii ye chlorocatechol degradation). Zvakataurwa kuti majini ari pa ICE anogona kutamiswa nekubatana ne frequency yakaderera kwazvo yekufambisa (10-8), nokudaro achiendesa zvinhu zvekuora kune anogamuchira (Basu na Phale, 2008; Phale et al., 2019).
Majini mazhinji anokonzera kuora kwecarbaryl ari pamaplasmid. Arthrobacter sp. RC100 ine maplasmid matatu (pRC1, pRC2 uye pRC300) ayo maviri maplasmid anobatana, pRC1 uye pRC2, anoronga maenzymes anoshandura carbaryl kuita gentisate. Kune rumwe rutivi, maenzymes anobatanidzwa mukushandurwa kwegentisate kuita central carbon metabolites ari pachromosome (Hayaatsu et al., 1999). Mabhakitiriya erudzi rweRhizobium. Strain AC100, inoshandiswa kushandurwa kwecarbaryl kuita 1-naphthol, ine plasmid pAC200, inotakura cehA gene inokochekera CH sechikamu cheTnceh transposon yakakomberedzwa neinsertion element-like sequences (istA uye istB) (Hashimoto et al., 2002). MuSphingomonas strain CF06, gene recarbaryl degradation rinofungidzirwa kuti riripo mumaplasmid mashanu: pCF01, pCF02, pCF03, pCF04, uye pCF05. Kubatana kweDNA kwemaplasmid aya kwakakwira, zvichiratidza kuvapo kwechiitiko chekudzokororwa kwemajini (Feng et al., 1997). Mu carbaryl-degrading symbiont inoumbwa nemarudzi maviri ePseudomonas, strain 50581 ine conjugative plasmid pCD1 (50 kb) inokodha mcd carbaryl hydrolase gene, nepo conjugative plasmid iri mustrain 50552 inokodha enzyme ye1-naphthol-degrading (Chapalamadugu naChaudhry, 1991). MuAchromobacter strain WM111, mcd furadan hydrolase gene iri pa100 kb plasmid (pPDL11). Jini iri rakaratidzwa kuti riripo pamaplasmid akasiyana (100, 105, 115 kana 124 kb) mumabhakitiriya akasiyana kubva kunzvimbo dzakasiyana dzenzvimbo (Parekh et al., 1995). MuPseudomonas sp. C5pp, majini ese anokonzera kuora kwecarbaryl ari mugenome inosvika 76.3 kb ye sequence (Trivesi et al., 2016). Kuongororwa kwemajini (6.15 Mb) kwakaratidza kuvapo kwemaMGE makumi mana nemaviri nemaGEI makumi matatu nematanhatu, ayo maMGE gumi nemanomwe aive mu supercontig A (76.3 kb) ane avhareji yeG+C content isina kuenzana (54–60 mol%), zvichiratidza kuti zviitiko zvekuchinjana kwemajini zvakatwasuka zvinogona kuitika (Trivesi et al., 2016). P. putida XWY-1 inoratidza kurongeka kwakafanana kwemajini anoparadza carbaryl, asi majini aya ari paplasmid (Zhu et al., 2019).
Pamusoro pekushanda zvakanaka kwemetabolism pamazinga ebiochemical uye genomic, tupukanana tunoratidzawo zvimwe zvinhu kana mhinduro dzakadai sechemotaxis, maitiro ekugadzirisa masero, kupatsanurwa kwezvikamu, kushandiswa kwakasarudzwa, kugadzirwa kwebiosurfactant, nezvimwewo, izvo zvinovabatsira kugadzirisa zvinosvibisa zvinonhuhwirira zviri nani munzvimbo dzakasvibiswa (Mufananidzo 7).
Mufananidzo 7. Nzira dzakasiyana dzekugadzirisa masero dzebhakitiriya rinoparadza hydrocarbon rinonhuhwirira kuti riparadze zvinobudirira zvinhu zvinosvibisa zvekunze.
Mhinduro dzemakemikari dzinoonekwa sezvinhu zvinowedzera kuparara kwezvinosvibisa zvipenyu munzvimbo dzakasiyana-siyana dzakasvibiswa. (2002) dzakaratidza kuti chemotaxis yePseudomonas sp. G7 kuenda ku naphthalene yakawedzera mwero wekuora kwe naphthalene munzvimbo dzemvura. Mhando yemhando yemusango G7 yakaora naphthalene nekukurumidza kupfuura mhando yemutant isina chemotaxis. Protein yeNahY (538 amino acids ine membrane topology) yakawanikwa ichibatanidzwa nemajini emetacleavage paplasmid yeNAH7, uye sema transducers emakemikari, puroteni iyi inoita seinoshanda se chemoreceptor yekuora kwe naphthalene (Grimm naHarwood 1997). Imwe ongororo yakaitwa naHansel et al. (2009) yakaratidza kuti puroteni iyi ine chemotactic, asi mwero wayo wekuora wakakwira. (2011) yakaratidza mhinduro yePseudomonas (P. putida) kune gaseous naphthalene, uko kupararira kwegasi kwakaguma nekuyerera kwakasimba kwenaphthalene kumasero, izvo zvaidzora mhinduro yekemotactic yemasero. Vaongorori vakashandisa maitiro aya ekemotactic kugadzira hutachiona hwaizowedzera mwero wekuora. Zvidzidzo zvakaratidza kuti nzira dzekemosensory dzinodzorawo mamwe mabasa emasero akadai sekupatsanurwa kwemasero, kudzorwa kwemasero, uye kuumbwa kwebiofilm, nokudaro zvichibatsira kudzora mwero wekuora. Zvisinei, kushandisa hunhu uhwu (chemotaxis) kuti huparadze zvinobudirira kunokanganiswa nezvikonzero zvakasiyana-siyana. Zvipingamupinyi zvikuru ndeizvi: (a) ma-paralogous receptors akasiyana anoziva macompounds/ligands akafanana; (b) kuvapo kwema-alternative receptors, kureva, energieous tropism; (c) kusiyana kwakakosha kwezvikamu zvepfungwa zvemhuri imwe chete yema-receptor; uye (d) kushaikwa kweruzivo pamusoro pemapuroteni makuru e-bacterial sensor (Ortega et al., 2017; Martin-Mora et al., 2018). Dzimwe nguva, kuora kwe aromatic hydrocarbons kunoburitsa metabolites/intermediates dzakawanda, izvo zvinogona kunge zviri chemotactic kune rimwe boka remabhakitiriya asi zvichisemesa kune mamwe, zvichiwedzera kuoma maitiro acho. Kuti tizive kudyidzana kwema ligands (aromatic hydrocarbons) nema chemical receptors, takagadzira ma hybrid sensor proteins (PcaY, McfR, naNahY) nekubatanidza sensor uye signaling domains dzePseudomonas putida naEscherichia coli, dzinonanga ma receptors e aromatic acids, TCA intermediates, uye naphthalene, zvichiteerana (Luu et al., 2019).
Pasi pesimba re naphthalene nedzimwe polycyclic aromatic hydrocarbons (PAHs), chimiro che bacterial membrane uye kusimba kwe microorganisms zvinochinja zvakanyanya. Zvidzidzo zvakaratidza kuti naphthalene inokanganisa kudyidzana kwe acyl chain kuburikidza ne hydrophobic interactions, nokudaro ichiwedzera kuzvimba uye fluidity ye membrane (Sikkema et al., 1995). Kuti irwise mhedzisiro iyi yakaipa, mabhakitiriya anodzora fluidity ye membrane nekushandura ratio uye fatty acid composition pakati pe iso/anteiso branched-chain fatty acids uye isomerizing cis-unsaturated fatty acids kuita trans-isomers inoenderana (Heipieper na de Bont, 1994). MuPseudomonas stutzeri inorimwa pakurapa naphthalene, saturated to unsaturated fatty acid ratio yakawedzera kubva pa1.1 kusvika pa2.1, nepo muPseudomonas JS150 ratio iyi yakawedzera kubva pa7.5 kusvika pa12.0 (Mrozik et al., 2004). Payakakura pa naphthalene, masero eAchromobacter KAs 3-5 akaratidza kuunganidzwa kwemasero akakomberedza makristaro e naphthalene uye kudzikira kwesimba repamusoro pemasero (kubva -22.5 kusvika -2.5 mV) pamwe chete ne cytoplasmic condensation uye vacuolization, zvichiratidza shanduko muchimiro chemasero uye hunhu hwepamusoro pemasero (Mohapatra et al., 2019). Kunyangwe shanduko dzemasero/pamusoro dzakabatana zvakananga nekutorwa zviri nani kwezvinosvibisa zvinonhuhwirira, nzira dzakakodzera dze bioengineering hadzina kunyatsogadziriswa. Kugadziriswa kwechimiro chemasero hakuna kushandiswa zvakanyanya kugadzirisa maitiro ehupenyu (Volke naNikel, 2018). Kubviswa kwemajini anokanganisa kupatsanurwa kwemasero kunokonzera shanduko muchimiro chemasero. Kubviswa kwemajini anokanganisa kupatsanurwa kwemasero kunokonzera shanduko muchimiro chemasero. MuBacillus subtilis, protein ye septum yemasero SepF yakaratidzwa kuti ine chekuita nekuumbwa kwe septum uye inodiwa pamatanho anotevera ekupatsanurwa kwemasero, asi haisi jini rinokosha. Kubviswa kwemajini anogadzira peptide glycan hydrolases muBacillus subtilis kwakaita kuti masero arebe, kuwedzera kwekukura, uye kugona kwekugadzira ma enzyme kwavandudzike (Cui et al., 2018).
Kugadziriswa kwenzira yekuparadza carbaryl kwakaitwa kuti kubudirire kuderedza Pseudomonas strains C5pp naC7 (Kamini et al., 2018). Zvinokurudzirwa kuti carbaryl iendeswe munzvimbo yeperiplasmic kuburikidza ne outer membrane septum uye/kana kuburikidza ne diffusible porins. CH i periplasmic enzyme inokonzeresa hydrolysis ye carbaryl kuenda ku 1-naphthol, iyo yakagadzikana, isingade mvura uye ine chepfu. CH inowanikwa mu periplasm uye ine affinity yakaderera ye carbaryl, nokudaro ichidzora kuumbwa kwe 1-naphthol, nokudaro ichidzivirira kuungana kwayo mumasero uye kuderedza chepfu yayo kumasero (Kamini et al., 2018). 1-naphthol inobuda inotakurwa kuenda ku cytoplasm kuburikidza ne inner membrane nekuparadzanisa uye/kana kupararira, uye yozosanganiswa kuita 1,2-dihydroxynaphthalene ne high-affinity enzyme 1NH kuti iwedzere kushandiswa mu central carbon pathway.
Kunyangwe tupukanana tune hunyanzvi hwemajini nemetabolism hwekukanganisa manyuko ekabhoni ye xenobiotic, chimiro chehierarchical chekushandiswa kwatwo (kureva, kushandiswa kwakasarudzika kwemasuko ekabhoni ari nyore pane akaomarara) chipingamupinyi chikuru pakuora kwehupenyu. Kuvapo nekushandiswa kwemasuko ekabhoni ari nyore kunoderedza majini anoisa ma enzymes anoparadza manyuko ekabhoni akaomarara/asingafarirwi akadai sePAHs. Muenzaniso wakanyatsodzidzwa ndewekuti kana glucose ne lactose zvikadyiswa pamwe chete kuEscherichia coli, glucose inoshandiswa zvakanyanya kupfuura lactose (Jacob naMonod, 1965). Pseudomonas yakashumwa kuti inoshatisa maPAH akasiyana-siyana uye xenobiotic compounds semanyuko ekabhoni. Huwandu hwekushandiswa kwemasuko ekabhoni muPseudomonas i organic acids > glucose > aromatic compounds (Hylemon naPhibbs, 1972; Collier et al., 1996). Zvisinei, pane musiyano. Zvinofadza kuti, Pseudomonas sp. CSV86 inoratidza chimiro chakasiyana chehierarchical chinoshandisa aromatic hydrocarbons (benzoic acid, naphthalene, nezvimwewo) pane glucose uye chinobatanidza aromatic hydrocarbons ne organic acids (Basu et al., 2006). Mubhakitiriya iyi, majini ekuparadza nekutakura aromatic hydrocarbons haaderedzwe kunyangwe paine kabhoni yechipiri senge glucose kana organic acids. Payakakura muglucose uye aromatic hydrocarbons medium, zvakaonekwa kuti majini ekufambisa nekugadzira glucose akaderedzwa, aromatic hydrocarbons akashandiswa muchikamu chekutanga chelog, uye glucose yakashandiswa muchikamu chechipiri chelog (Basu et al., 2006; Choudhary et al., 2017). Kune rumwe rutivi, kuvapo kwe organic acids hakuna kukanganisa kuratidzwa kwe aromatic hydrocarbon metabolism, saka bhakitiriya iyi inotarisirwa kuva mhando ye biodegradation studies (Phale et al., 2020).
Zvinozivikanwa kuti hydrocarbon biotransformation inogona kukonzera oxidative stress uye kuwedzera kweantioxidant enzymes mu microorganisms. Kusashanda zvakanaka kwe naphthalene biodegradation mu stationary phase cells uye muine chepfu compounds kunotungamira pakuumbwa kwe reactive oxygen species (ROS) (Kang et al. 2006). Sezvo naphthalene-degrading enzymes ine iron-sulfur clusters, pasi pe oxidative stress, iron iri mu heme uye iron-sulfur proteins icha oxidized, zvichikonzera protein inactivation. Ferredoxin-NADP+ reductase (Fpr), pamwe chete ne superoxide dismutase (SOD), inobatanidza reversible redox reaction pakati peNADP+/NADPH nemamorekuru maviri eferredoxin kana flavodoxin, nokudaro ichibvisa ROS uye kudzoreredza iron-sulfur center pasi pe oxidative stress (Li et al. 2006). Zvakataurwa kuti Fpr neSodA (SOD) muPseudomonas zvinogona kukonzerwa nekushushikana kwe oxidative, uye kuwedzera kweSOD ne catalase activity kwakaonekwa mumhando ina dzePseudomonas (O1, W1, As1, uye G1) panguva yekukura pasi pemamiriro ekuwedzera naphthalene (Kang et al., 2006). Zvidzidzo zvakaratidza kuti kuwedzerwa kwe antioxidants senge ascorbic acid kana ferrous iron (Fe2+) kunogona kuwedzera kukura kwe naphthalene. Rhodococcus erythropolis payakakura mu naphthalene medium, kunyorwa kwemajini e oxidative stress-related cytochrome P450 anosanganisira sodA (Fe/Mn superoxide dismutase), sodC (Cu/Zn superoxide dismutase), uye recA kwakawedzerwa (Sazykin et al., 2019). Kuenzanisa uwandu hweproteomic ongororo yemasero ePseudomonas akakurira mu naphthalene kwakaratidza kuti kuwedzera kwemapuroteni akasiyana-siyana ane chekuita nemhinduro yekushushikana kwe oxidative inzira yekugadzirisa kushushikana (Herbst et al., 2013).
Zvipukanana zvakanzi zvinogadzira biosurfactants pasi pesimba rezvinhu zvinogadzira kabhoni inonyungudika mumvura. Izvi zvinogadzira amphiphilic surface active compounds zvinogona kugadzira ma aggregates pamafuta-mvura kana mhepo-mvura. Izvi zvinokurudzira pseudo-solubilization uye zvinoita kuti kunyungudika kwemahydrocarbon anonhuhwirira kuwedzere, zvichikonzera kuora kwebiogradation kunobudirira (Rahman et al., 2002). Nekuda kwehunhu uhu, biosurfactants dzinoshandiswa zvakanyanya mumaindasitiri akasiyana-siyana. Kuwedzera makemikari esurfactants kana biosurfactants kumabhakitiriya kunogona kuwedzera kushanda zvakanaka uye mwero wekuora kwehydrocarbon. Pakati pezvinhu zvinogadzira biosurfactants, rhamnolipids inogadzirwa nePseudomonas aeruginosa yakaongororwa zvakanyanya uye yakatsanangurwa (Hisatsuka et al., 1971; Rahman et al., 2002). Pamusoro pezvo, mamwe marudzi e biosurfactants anosanganisira lipopeptides (mucins kubva kuPseudomonas fluorescens), emulsifier 378 (kubva kuPseudomonas fluorescens) (Rosenberg naRon, 1999), trehalose disaccharide lipids kubva kuRhodococcus (Ramdahl, 1985), lichenin kubva kuBacillus (Saraswathy naHallberg, 2002), uye surfactant kubva kuBacillus subtilis (Siegmund naWagner, 1991) uye Bacillus amyloliquefaciens (Zhi et al., 2017). Aya ma surfactants ane simba akaratidzwa kuti anoderedza kushushikana kwepamusoro kubva pa72 dynes/cm kusvika pasi pe30 dynes/cm, zvichibvumira kunyudzwa kuri nani kwehydrocarbon. Zvakataurwa kuti Pseudomonas, Bacillus, Rhodococcus, Burkholderia nedzimwe mhando dzebhakitiriya dzinogona kugadzira mhando dzakasiyana dze rhamnolipid ne glycolipid-based biosurfactants kana dzikakura mu naphthalene ne methylnaphthalene media (Kanga et al., 1997; Puntus et al., 2005). Pseudomonas maltophilia CSV89 inogona kugadzira extracellular biosurfactant Biosur-Pm kana ikakura pa aromatic compounds dzakadai se naphthoic acid (Phale et al., 1995). Kinetics yekuumbwa kweBiosur-Pm yakaratidza kuti kugadzirwa kwayo kunoenderana nekukura uye pH. Zvakaonekwa kuti huwandu hweBiosur-Pm hunogadzirwa nemasero pa neutral pH hwakanga hwakakwira kupfuura hwepH 8.5. Masero akakura pa pH 8.5 aive ne hydrophobic zvakanyanya uye aive ne affinity yakakura ye aromatic uye aliphatic compounds kupfuura masero akakura pa pH 7.0. MuRhodococcus spp. N6, huwandu hwakakwira hwecarbon kuenda kunitrogen (C:N) uye kushaya iron ndizvo zvinhu zvakanaka pakugadzirwa kwe extracellular biosurfactants (Mutalik et al., 2008). Kuedza kwakaitwa kuvandudza biosynthesis ye biosurfactants (surfactins) nekuvandudza strains uye fermentation. Zvisinei, titer ye surfactant mu culture medium yakaderera (1.0 g/L), izvo zvinopa dambudziko pakugadzirwa kukuru (Jiao et al., 2017; Wu et al., 2019). Nokudaro, nzira dze genetic engineering dzakashandiswa kuvandudza biosynthesis yayo. Zvisinei, kugadziriswa kwayo kweinjiniya kwakaoma nekuda kwehukuru hwe operon (∼25 kb) uye kurongeka kwakaoma kwe biosynthetic ye quorum sensing system (Jiao et al., 2017; Wu et al., 2019). Kugadziriswa kwakasiyana-siyana kwemajini kwakaitwa mumabhakitiriya eBacillus, kunyanya kunangana nekuwedzera kugadzirwa kwesurfactin nekutsiva promoter (srfA operon), kuwedzera huwandu hweprotein yesurfactin export YerP uye zvinhu zvinodzora ComX naPhrC (Jiao et al., 2017). Zvisinei, nzira idzi dzekugadzira majini dzakangogadzirisa majini kamwe chete kana mashoma uye hadzisati dzasvika pakugadzirwa kwekutengeserana. Nokudaro, kudzidza kwakawanda kwenzira dzekugadzirisa ruzivo kwakakosha.
Zvidzidzo zvekuora kwePAH zvinonyanya kuitwa pasi pemamiriro ezvinhu akajairwa murabhoritari. Zvisinei, panzvimbo dzakasvibiswa kana munzvimbo dzakasvibiswa, zvinhu zvakawanda zvisingakuvadzi zvipenyu (tembiricha, pH, okisijeni, kuwanikwa kwezvinovaka muviri, kuwanikwa kwesubstrate, mamwe ma xenobiotics, kudziviswa kwekugadzirwa kwemichero, nezvimwewo) zvakaratidzwa kuti zvinochinja uye zvinokanganisa kugona kwehutachiona kuora.
Kupisa kune simba guru pakuora kwePAH. Sezvo tembiricha ichiwedzera, huwandu hweokisijeni yakanyungudutswa hunoderera, izvo zvinokanganisa metabolism ye aerobic microorganisms, sezvo ichida molecular oxygen seimwe ye substrates dze oxygenases dzinoita hydroxylation kana ring cleavage reactions. Zvinowanzo cherechedzwa kuti kupisa kwakanyanya kunoshandura maPAH emubereki kuita makemikari ane chepfu, nokudaro zvichidzivisa kuora kwebiogradation (Muller et al., 1998).
Zvakaonekwa kuti nzvimbo zhinji dzine tsvina yePAH dzine pH yakawandisa, dzakadai senzvimbo dzine acid mine drainage dzakasvibiswa (pH 1–4) uye nzvimbo dzegasi rechisikigo/marasha dzakasvibiswa nealkaline leachate (pH 8–12). Mamiriro aya anogona kukanganisa zvakanyanya maitiro ekuora kwezvipenyu. Nokudaro, usati washandisa tupukanana kuti ugadzire makemikari, zvinokurudzirwa kugadzirisa pH nekuwedzera makemikari akakodzera (ane mukana wekuderedza oxidation pakati nepakati kusvika pasi) akadai seammonium sulfate kana ammonium nitrate yealkaline soils kana liming necalcium carbonate kana magnesium carbonate yenzvimbo dzine acidic (Bowlen et al. 1995; Gupta naSar 2020).
Kuwanikwa kweokisijeni kunzvimbo yakakanganiswa ndicho chinhu chinotadzisa PAH kuora kwemvura. Nekuda kwemamiriro e redox enzvimbo, maitiro ekugadzirisa okisijeni munzvimbo iyi anowanzoda kuiswa kweokisijeni kubva kunze (kurima, kuparadzira mweya, uye kuwedzerwa kwemakemikari) (Pardieck et al., 1992). Odenkranz et al. (1996) vakaratidza kuti kuwedzera magnesium peroxide (chinhu chinoburitsa okisijeni) kune aquifer yakasviba kunogona kugadzirisa BTEX compounds zvinobudirira. Imwe ongororo yakaongorora kuora kwe phenol neBTEX munzvimbo iyi mu aquifer yakasviba nekuisa sodium nitrate nekuvaka matsime ekuburitsa okisijeni kuti pave nekugadzirisa okisijeni zvinobudirira (Bewley naWebb, 2001).