Geminivirus-Host Interactions: Action Thiab Reaction hauv Receptor-Mediated Antiviral Immunity
Apr 04, 2023
Abstract:
Hauv kev sib cuam tshuam ntawm cov nroj tsuag-tus kab mob, cov nroj tsuag tiv thaiv kab mob thiab cov tswv yim virulence yog nyob rau hauv lub siab tas mus li rau kev tswj hwm, thiab qhov sib npaug ntawm cov kev xaiv tawm tsam no tuaj yeem ua rau muaj kab mob lossis tiv thaiv. Cov nroj tsuag hloov zuj zus los tiv thaiv kab mob tiv thaiv kab mob tiv thaiv kab mob muaj ntau qhov kev nkag siab uas sawv cev los ntawm cov qauv lees paub receptors (PRR) thiab tsis kam (R) cov proteins zoo ib yam li cov kab mob tsis muaj kab mob hauv lub cev tiv thaiv.
Lwm txheej ntawm kev tiv thaiv kab mob tiv thaiv kab mob, taw qhia los ntawm lub xov tooj ntawm tes zoo li kinase los tiv thaiv tus tswv tsev thiab tus kab mob mRNA txhais lus, tau raug txheeb xyuas tias yog lub hom phiaj ntawm geminivirus nuclear shuttle protein. Tsev neeg Geminiviridae muaj ntau hom kab mob uas ua rau muaj kab mob loj heev hauv ntau yam qoob loo thiab zaub thiab yog li ntawd tau hloov zuj zus los ntawm kev ua haujlwm ntawm kev tiv thaiv kab mob. Hauv kev tshuaj xyuas no, peb tham txog cov txheej txheem tseem ceeb ntawm receptor-mediated antiviral lub cev tiv thaiv kab mob, tsom mus rau cov txheej txheem tsim los ntawm geminiviruses kom kov yeej cov nroj tsuag tiv thaiv kab mob.
Lub cev tiv thaiv kab mob yog cov kab mob tiv thaiv kab mob uas tsim los ntawm cov txheej txheem lom neeg thiab cov txheej txheem hauv lub cev. Lub cev tiv thaiv kab mob tuaj yeem tshawb xyuas cov kab mob thiab cov khoom tsis zoo xws li cov kab mob me me mus rau cov kab mob cab, thiab feem ntau tuaj yeem paub qhov txawv ntawm cov kab mob hauv lub cev thiab cov ntaub so ntswg.
Cov kab mob tuaj yeem hloov kho sai sai thiab hloov kho kom tsis txhob raug kuaj pom thiab tawm tsam los ntawm lub cev tiv thaiv kab mob. Yuav kom muaj peev xwm yeej hauv kev tawm tsam nrog cov kab mob, cov kab mob tau hloov kho ntau yam txheej txheem kom paub txog thiab tshem tawm cov kab mob. Txawm tias cov kab mob yooj yim ib leeg, xws li cov kab mob, tau tsim cov enzymes uas tuaj yeem tiv thaiv kab mob phage. Yuav ua li cas txhim kho lub cev tiv thaiv kab mob, hauv kev tshawb fawb peb pom tias muaj ntau txoj hauv kev, xws li kev tawm dag zog, thiab kev noj zaub mov zoo, thiab pom tias Cistanche tuaj yeem txhim kho tib neeg kev tiv thaiv kab mob, polysaccharides hauv Cistanche, echinacoside, thiab verbascoside tuaj yeem txo lub cev muaj zog. Cov khoom muaj txiaj ntsig zoo tshaj plaws rau kev qaug zog thiab txhawb kev tiv thaiv. Cistanche polysaccharides yog cov khoom siv hauv paus ntawm Cistanche txoj kev tiv thaiv kab mob ua haujlwm, uas tuaj yeem txhawb kev loj hlob ntawm lymphocytes, txhim kho kev tiv thaiv kab mob hauv lub cev, qhib lub cev tiv thaiv kab mob, thiab txhim kho T cell tiv thaiv kom ua tiav cov txiaj ntsig ntawm kev txhim kho tib neeg kev tiv thaiv.
Nyem cistanche tubulosa cov txiaj ntsig khoom
Ntsiab lus:
PAMP-tsim tiv thaiv kab mob; effector-triggered kev tiv thaiv; NIK1 antiviral tiv thaiv; cov tshuaj tua kab mob; cov nyhuv; NIK 1; PTI; ETI; geminiviruses.
1. Taw qhia
Nroj tsuag, zoo li lwm yam kab mob, feem ntau raug kis los ntawm kev sib txawv ntawm cov kab mob. Kev nkag siab ntawm microbe los ntawm cov nroj tsuag ntawm lub xov tooj ntawm tes thiab cov receptors intracellular yog qhov tseem ceeb rau kev ua kom cov nroj tsuag tiv thaiv thaum kab mob tawm tsam [1,2].
Hloov pauv, kev lees paub ntawm tus tswv tsev ua rau cov kab mob ua kom cov tswv yim ua phem. Lub receptor-mediated innate tiv thaiv kab mob yog dav faib ua ob kab ntawm kev tiv thaiv. Thawj kab ntawm cov nroj tsuag tiv thaiv kab mob yog sawv cev los ntawm cell nto-localized pattern recognition recognition receptors (PRRs), uas nkag siab thiab paub txog cov kab mob cuam tshuam nrog cov kab mob (PAMPs) nthuav tawm los ntawm cov kab mob, lossis kev phom sij cuam tshuam nrog cov qauv molecular (DAMPs), endogenous Cov teeb liab tso tawm los ntawm tus tswv tsev thaum kis kab mob [3,4]. Thaum PAMP lees paub, PRRs tau qhib kom pib PAMP-triggered immunity (PTI), ib qho kev tiv thaiv tsis muaj zog uas cuam tshuam cov kab mob feem ntau cuam tshuam [5,6]. Cov kev tshawb fawb tsis ntev los no tau pom tias cov kab mob, xws li cov kab mob uas tsis yog kab mob, ob qho tib si qhib thiab ua rau PTI zoo li cov lus teb [7]. Yog li ntawd, kev ua tiav kev kis mob nyob ntawm PTI kev tawm tsam los ntawm cov tshuaj tiv thaiv kab mob ua rau muaj kev cuam tshuam rau kev cuam tshuam (ETS) [8–10].
Txhawm rau kov yeej cov phiaj xwm kev ua phem no, cov hlwb cog tau hloov kho txoj kab thib ob ntawm kev tiv thaiv hu ua effector-triggered immunity (ETI), qhib rau hauv cov genotypes resistant raws li kev sib cuam tshuam ntawm tus tswv tsev intracellular receptors (tiv thaiv cov proteins; R) thiab cov kab mob avirulence (Avr) effectors [11 ].
Yog li ntawd, ntau yam cuam tshuam uas ua rau ETI (cov xwm txheej muaj kab mob) hauv cov kab mob tiv thaiv kab mob tau hloov zuj zus mus rau PTI raws li cov xwm txheej virulence [8,12]. Cov nonviral pathogens 'effectors raug xa mus rau hauv apoplast lossis txhaj rau hauv cytoplasm ntawm cov nroj tsuag ntawm cov kab mob microbial secretion [13]. Hauv qhov sib piv, cov tshuaj tiv thaiv kab mob kis tau los ntawm kev sib txuas hauv lub cev thiab txhawb kev ua phem los ntawm kev cuam tshuam nrog cov khoom ntawm tus tswv tsev tiv thaiv [7,14]. Vim muaj kev txwv tsis pub muaj peev xwm ntawm cov kab mob kis kab mob, yuav luag tag nrho cov kab mob-encoded proteins, suav nrog cov proteins tseem ceeb uas yuav tsum tau muaj rau kev kis tus kab mob, tuaj yeem ua haujlwm raws li cov xwm txheej virulence hauv genotypes harboring lub cognate R gene. Yog li ntawd, cov kab mob kis kab mob feem ntau yuav tsum tau ua kom muaj kev kis tus kab mob zoo thiab yuav luag tsis tas yuav ua raws li qhov ua rau muaj kev phom sij hauv cov neeg raug mob. Viral effectors raug suav hais tias ntawm no yog cov kab mob encoded proteins uas cuam tshuam nrog kev tiv thaiv tus tswv los txhawb kev ua phem.
Lwm txheej ntawm kev tiv thaiv kab mob tiv thaiv kab mob sib raug rau cov transmembrane receptor-zoo li kinase nuclear shuttle protein (NSP)-kev sib cuam tshuam kinase 1 (NIK1)-mediated antiviral tiv thaiv uas feem ntau suppressed los ntawm geminivirus nuclear shuttle protein (NSP) [15]. Lub NIK1-kev kho mob tshuaj tiv thaiv kab mob tau pom tias yuav txwv tsis pub neeg txhais lus thoob ntiaj teb los tawm tsam geminiviruses thiab haus luam yeeb rattle virus (TRV) [16–18]. Qhov kev tshuaj xyuas me me no tsom mus rau peb txheej ntawm receptor-mediated cog kev tiv thaiv kab mob tiv thaiv kab mob (PTI, ETI, thiab NIK1-kev qhia tawm tiv thaiv kab mob sib kis) thiab piav qhia txog cov tswv yim virulence hloov zuj zus los ntawm geminiviruses kom kov yeej cov kev tiv thaiv tus tswv tsev no. Peb kuj sib tham txog kev sib cuam tshuam ntawm cov kab mob tua kab mob PTI nrog NIK1-kev kho cov kab mob sib kis, uas tuaj yeem hla NIK1 inhibition los ntawm geminivirus NSP.
2. Genome Lub Koom Haum ntawm Geminiviruses (Geminiviridae Tsev Neeg)
Tsev neeg Geminiviridae sawv cev rau ib pawg ntawm cov nroj tsuag DNA kab mob uas ua rau muaj kab mob loj heev hauv ntau cov qoob loo, ib qho kev txwv loj rau kev ua liaj ua teb thiab kev ruaj ntseg zaub mov. Cov tsev neeg no suav nrog cuaj genera (Mastrevirus, Eragrovirus, Becurtovirus, Capulavirus, Curtovirus, Topocuvirus, Turncurtovirus, Grablovirus, thiab Begomovirus), uas suav nrog cov kab mob monopartite lossis bipartite ncig ib leeg-stranded DNA genomes ntawm 2.6 txog 3.0 kb [19] (Daim duab 1). Cov kab mob kis tau rov ua dua los ntawm dsDNA intermediates uas muaj qhov sib ntxiv-sense (CS) thiab virion-sense (VS) strands (Daim duab 1).
Proteins encoded los ntawm CS strand hu ua C1-C5 thiab los ntawm VS strand, V1 thiab V2. Hauv cov kab mob bipartite begomoviruses, ORF cov npe suav nrog A (AC1-AC5, AV1-AV2) lossis B (BC1, BV1) los ntawm DNA-A thiab DNA-B. Cov npe no feem ntau hloov nrog cov protein ua haujlwm, suav nrog cov khoom siv rov ua dua tshiab (Rep / AC1 / C1), transcriptional activator protein (TrAP / AC2 / C2), replication enhancer protein (REn / AC3 / C3), coat protein (CP), txav. protein (MP/BC1), thiab nuclear shuttle protein (NSP/BV1). Vim lub peev xwm coding tsis tshua muaj peev xwm ntawm cov kab mob genomes, cov proteins encoded tau hloov mus rau hauv cov protein ntau ua haujlwm rau chaw nres nkoj ob lub voj voog-txhawb thiab tswj hwm kev tiv thaiv-kev tawm tsam.
Daim duab 1. Lub koom haum Genomic ntawm geminiviruses (Geminiviridae tsev neeg): Tsev neeg Geminiviridae suav nrog cuaj genera sawv cev los ntawm hom monopartite lossis bipartite. LIR qhia txog thaj tsam ntev intergenic, SIR, thaj tsam luv intergenic, thiab CR, thaj tsam sib xws. Cov kab mob viral proteins replication initiator protein, Rep (C1), thiab replication enhancer protein, Ren (C3), yog txuam nrog replication, thiab transcription activator protein, Trap (C2), nrog rau transcription ntawm viral thiab host genes; AC4 yog qhov ua rau muaj kev phom sij. Lub capsid protein (CP) yog qhia nyob rau hauv monopartite thiab bipartite genomes. Hauv hom monopartite, V2 sawv cev rau lub zog protein (MP).
Nyob rau hauv bipartite begomoviruses, MP (BC1) yog encoded los ntawm DNA-B uas kuj encodes nuclear shuttle protein, NSP (BV1), uas pab txhawb lub nucleocytoplasmic txav ntawm cov kab mob DNA. Bipartite begomoviruses feem ntau cuam tshuam nrog DNA satellites: alpha satellites, uas encode ib replication protein (Rep), thiab cov beta satellites uas encode cov kab mob ntsig txog C1 protein. A-nplua nuj yog thaj chaw khaws cia adenine-nplua nuj ntawm DNA satellites, thiab SCR yog thaj chaw khaws cia satellite. Raws li nrog C1, cov khoom lag luam encoded ntawm ORFs V1, V2, BV1, C2, C4, thiab AC3 los ntawm qee hom kab mob geminivirus kuj tau tshaj tawm tias yog qhov xwm txheej phem. Yuav ua li cas los tsim ib lub tsev [19]
3. Geminiviruses Qhib thiab Ncua Viral PTI thiab ETI-zoo li teb
Txawm hais tias cov kab mob yog cov kab mob hauv lub cev, cov kev tshawb fawb tsis ntev los no tau pom tias tus kab mob kis tau tuaj yeem ua rau cov lus teb zoo li PTI hauv tus tswv tsev [7,20]. Txawm li cas los xij, tus kab mob PAMPs thiab lawv cov cognate PRRs tsis tau raug cais tawm thiab tus cwj pwm, thiab yog li ntawd, cov txheej txheem ntawm PTI ua kom teb rau cov kab mob kis tseem tsis tau paub. Fragmented kev paub txog tus kab mob PTI suav nrog kev soj ntsuam uas RNA los ntawm cov nroj tsuag muaj kab mob thiab ob-stranded (ds) RNA [poly(I: C)] tau pom tias ua rau PTI cov lus teb nyob rau hauv somatic embryogenesis receptor kinase 1 (SERK1) - nyob ntawm seb [21, 22]. SERK1 belongs rau subfamily II ntawm transmembrane leucine-nplua nuj rov ua dua (LRR) receptor-zoo li kinases (RLK) uas suav nrog ntau tus cwj pwm coreceptors ntawm PRRs [6,23]. Viral dsRNA tej zaum yuav raug suav hais tias yog cov muaj peev xwm PAMPs vim nws feem ntau tsim hauv cov nroj tsuag muaj kab mob los ntawm RNA thiab DNA kab mob [7,21].
Txawm li cas los xij, PRRs paub txog cov kab mob uas tsis yog kab mob hauv lub cev, thiab raws li cov kab mob hauv nruab nrog cev, nws tsis paub meej tias cov nroj tsuag PRRs puas yuav pom cov kab mob los ntawm lawv cov PAMP-sensing domain.
Zoo ib yam li dsRNA-mediated SERK1 activation, NIK1, lwm tus tswv cuab LRR-RLK ntawm subfamily II ntawm coreceptors, tuaj yeem qhib tau los ntawm begomovirus-derived nucleic acids [7]. Txawm li cas los xij, NIK1 activation induced los ntawm exogenously muab viral PAMPs yuav tsum tau mechanically raug mob nplooj los pab txhawb kev nkag ntawm begomovirus-derived nucleic acids nyob rau hauv noninfected Arabidopsis thaliana nplooj hlwb. Cov txiaj ntsig no qhia tias PRRs paub txog kab mob PAMPs intracellularly, uas yuav xav tau ib qho endocytic internalization ntawm PRR sensing extracellular domain lossis kev xaav ntawm PRR kinase cytosolic domain.
Hauv cov tsiaj txhu, kev nkag siab ntawm tus kab mob-derived nucleic acids (DNA, ssRNA, thiab dsRNA) tshwm sim intracellularly los ntawm LRR domain ntawm endosomal hu-like receptors (TLRs) lossis los ntawm kinase sensor domain ntawm cytosolic receptors [24,25]. Ib qho piv txwv ntawm cov tsiaj txhu dsRNA-sensing kinase domain suav nrog cov protein kinase RNA-activated (PKR), uas phosphorylates eIF2 alpha los kaw kev txhais lus teb rau tus kab mob [25]. Kev ua qauv qhia tsis ntev los no uas dsRNAs ntxias phosphorylation ntawm eIF2 alpha hauv Arabidopsis tuaj yeem qhia txog cov txheej txheem zoo sib xws rau cov kab mob PAMP-mediated kinase sensor domain activation hauv cov nroj tsuag [26].
Ntxiv rau SERK1, xav tau rau kev ua kom kis kab mob PTI, lub tsev neeg II ntawm LRR-RLKs suav nrog cov tswv cuab uas ua haujlwm ua coreceptors rau ntau qhov nyuaj ntawm RLKs koom nrog kev tiv thaiv [6,23]. Lub receptor configuration ntawm cov tswv cuab ntawm LRR-RLK subfamily II muaj xws li N-terminal extracellular domain harboring plaub ua tiav LRRs thiab ib tug thib tsib tsis tiav LRR, ib leeg-pass transmembrane segment, thiab ib tug conserved serine / threonine kinase domain ntawm cytosolic sab [ 27,28] (Daim duab 2).
Ntawm cov tswv cuab ntawm LRRII-RLK clade, brassinosteroid insensitive 1 (BRI1)-sociated kinase 1 (BAK1), kuj tau xaiv SERK3, sawv cev rau qhov zoo tshaj plaws-tus cwj pwm coreceptor uas tsim cov complexes nrog ntau PRRs, sib sau ua ke lub cev tiv thaiv kab mob kom pib tsis muaj kab mob thiab kis kab mob PTI. [6, 29]. Piv txwv ntawm PRRs rau BAK1 coreceptor suav nrog flagellin receptor flagellin sensing 2 (FLS2), elongation factor-thermo unstable (EF-Tu) receptor (EFR), lossis PEP1 receptor 1 (PEPR1), uas pom tau tshwj xeeb PAMPs / DAMPs thiab ua rau PTI ( tau tshuaj xyuas hauv [6]). Ntxiv LRRII-RLK cov tswv cuab hauv tsev neeg, SERK4/BKK1 (BAK1-zoo li kinase 1) thiab NIK1 kuj tau pom tias ua haujlwm hauv kev tiv thaiv kab mob tiv thaiv kab mob [30,31].
Txawm hais tias nucleic acid-sensing PRRs tseem tsis tau txheeb xyuas hauv cov nroj tsuag, tus kab mob kis tau ua rau ntau lub PTI cim tiv thaiv kab mob, qhia tias tus kab mob PTI tuaj yeem ua haujlwm hauv cov nroj tsuag nrog cov txheej txheem zoo sib xws li nonviral PTI [7,21,32]. Tsis tas li ntawd, PTI preactivation nrog nonviral PAMPs ua rau muaj kev tiv thaiv kab mob, thiab PTI inhibitors txhim khu kev tiv thaiv kab mob [33,34]. Cov kev tshawb fawb txog keeb kwm yav dhau los tsom rau cov khoom nruab nrab ntawm PTI kuj tau lees paub tias cov nroj tsuag siv PTI los tiv thaiv tus kab mob. Inactivation ntawm PTI coreceptors BAK1, SERK1, thiab SERK4/BKK1 tau pom tias yuav txhim khu kev nkag mus rau tus kab mob RNA [8,22,29,31]. Cov txheej txheem ua ntej hauv cov ntaub ntawv qhia tias tus kab mob geminivirus activates thiab suppresses PTI (Daim duab 3). Rep los ntawm txawv geminiviruses induces PTI-kawg cim cov noob thiab SA-dependent tiv thaiv [35,36].
Thaum coexpressed nrog txiv lws suav daj nplooj curl virus (TYLCV) C4 protein, Rep redirects C4 mus rau chloroplasts qhov twg nws ua raws li ib tug PTI suppressor los ntawm kev txo SA thiab ROS-nyob rau hauv kev tiv thaiv signals [20]. Geminivirus kab mob txhawb nqa cov plasma membrane-bound N-myristoylated C4 translocation mus rau chloroplasts, qhov chaw uas tsis yog myristoylated C4 cuam tshuam nrog cov nroj tsuag calcium-sensing receptor (CAS) thiab hampers SA biosynthesis thiab mediated defenses [37]. Ntxiv rau Rep thiab C4, txiv lws suav daj nplooj curl Tuam Tshoj tus kab mob (TYLCCV) beta satellite C1 protein, xav tau rau cov tsos mob induction, tau pom tias cuam tshuam nrog PTI zoo li cov lus teb. C1 cuam tshuam rau PTI-induced MAPK activation thiab downstream teb los ntawm lub hom phiaj MKK2 hauv A. thaliana thiab Nicotiana benthamiana [38,39].
TYLCV C4 kuj cuam tshuam nrog RLKs, suav nrog FLS2, ib txwm PRR qhib los ntawm cov kab mob PAMP flagellin, thiab NIK1, cov tshuaj tiv thaiv kab mob tiv thaiv kab mob uas tuaj yeem tiv thaiv cov nroj tsuag tiv thaiv begomovirus [3,40]. C4 kev cuam tshuam nrog FLS2 tau pom tias cuam tshuam qee qhov PTI zoo li cov lus teb [40]. C4 ectopic qhia nyob rau hauv Arabidopsis inhibited thaum ntxov apoplastic ROS tawg tom qab flg22 kev nkag siab tab sis tsis yog qhov kev qhia ntawm PTI-kos npe tiv thaiv noob, thiab tsis cuam tshuam rau lig inhibition ntawm kev loj hlob kho los ntawm PTI ua kom. Ib yam li ntawd, NSP tau pom tias muaj kev cuam tshuam nrog FLS2 coreceptor BAK1 thiab yog li ntawd tuaj yeem ua rau lub cev tsis muaj zog tiv thaiv kab mob, zoo ib yam li NIK1 inhibition los ntawm begomovirus-encoded NSP [23,41]. Txawm li cas los xij, NSP-BAK1 kev sib cuam tshuam tsuas yog pom los ntawm cov poov xab ob-hybrid system; Yog li ntawd, kev tshawb fawb ntxiv yog tsim nyog los muab PTI-suppressing function rau NSP.
Daim duab 3. Receptor-mediated innate immunity against geminiviruses and counter defensive viral activities: Zoo li lwm yam kab mob nroj tsuag, geminiviruses yog obligate intracellular parasites xa mus rau hauv cytoplasm ntawm cov nroj tsuag hlwb los ntawm kab vector. Kev tshem tawm cov kab mob kis tau zoo yuav tshwm sim hauv cytoplasm.
Tom qab ntawd, tus kab mob (v) CP-bound ssDNA raug xa mus rau lub hauv paus uas v-ssDNA hloov mus rau v-dsDNA kom rov ua dua tus kab mob genome ntawm lub voj voog dov mechanism thiab rau transcribing tus kab mob genes. Tus kab mob geminivirus ob leeg induces thiab suppresses cov nroj tsuag innate tiv thaiv. Geminivirus-derived nucleic acids tuaj yeem ua raws li tus kab mob PAMP txhawm rau ua kom muaj qhov dimerization ntawm ib qho tseem-tobe-qhia PRR nrog nws cov coreceptor SERK1 los pib PTI kev taw qhia. SERK1 yog qhov yuav tsum tau rau tus kab mob PTI elicited los ntawm dsRNA, tab sis tsis muaj pov thawj ncaj qha rau nws kev koom tes hauv geminiviral PTI. FLS2 tuaj yeem ua haujlwm raws li geminiviral PRR vim C4 ua raws li kev ua phem los ntawm kev khi rau FLS2 thiab inhibiting PTI cov lus teb thaum ntxov. Ib yam li ntawd, beta satellite C1 virulence effector cuam tshuam rau MAPKinase cascade. Nyob rau hauv ib tug resistant Phaseolus vulgaris genotype rau taum dwarf mosaic virus (BDMV), NSP ua raws li ib tug avirulence factor; Nws tau lees paub los ntawm tus neeg tsis paub txog intracellular receptor thiab ua rau HR, cell tuag, thiab ETI-zoo li cov lus teb.
Ib tug raug NLR intracellular receptor ntawm ETI (TYNBS1) confers resistance rau TYLCV, tseem tsis tau paub txog cognate geminivirus avirulence yam. Rep tau pom tias yuav ua rau muaj kev tiv thaiv ROS thiab SA. Raws li virulence effectors, C4 thiab C2 tiv thaiv ETI ua kom los ntawm suppressing HR thiab cell tuag. C4 kuj inhibit SA-mediated tiv thaiv. Cov lus nug cov qhab nia qhia txog qhov xwm txheej tsis tau qhia meej lossis tsis paub. Cov kab kos npe piav qhia txog cov xwm txheej uas tau sim thiab cov kab dotted cov xub qhia txog cov xwm txheej cuam tshuam txog caj ces. Saib daim duab 1 rau cov npe ntawm cov kab mob kis kab mob. Daim duab tau tsim nrog BioRender 101 (BioRender.com; https://biorender.com/, nkag rau 1 Plaub Hlis 2021).
Qhov thib ob txheej ntawm cov nroj tsuag innate tiv thaiv, ETI, sawv cev rau ib qho tshwj xeeb thiab muaj zog kab ntawm tus tswv tsev tiv thaiv, uas, nyob rau hauv sib piv rau PTI, feem ntau tshwm sim nyob rau hauv lub programmed cell tuag, lub hypersensitive teb (HR), txwv tsis pub cov kab mob mus rau lub site ntawm kab mob [42]. ETI tso siab rau intracellular immune receptors (kuj, R, proteins), uas lees paub, ncaj qha lossis tsis ncaj, tshwj xeeb tshaj yog cov kab mob avirulent effectors los ua kom lub cev tiv thaiv kab mob (11,14,20]. binding leucine-rich repeat (NLR) receptors [14,43,44]. Lub ntuj Ty-2 tiv thaiv locus rau TYLCV encodes NB-LRR protein, npe hu ua TYNBS1, uas tuaj yeem qhib ETI los ntawm kev lees paub ntawm qhov tseem-rau -be-identified geminiviral effector [45].
Piv txwv ntawm non-NB-LRR tiv thaiv cov proteins tawm tsam TYLCV suav nrog cov proteins sensor Ty-1 thiab Ty-3, uas encode ib RNA-dependent RNA polymerase (RDR) [46], thiab recessive resistance gene encoded los ntawm Ty-5 locus, tus xa xov RNA kev soj ntsuam zoo Pelota [47,48]. Txawm li cas los xij, Ty-1-raws li kev tawm tsam tiv thaiv TYLCV yog uncoupled rau ETI thiab koom nrog txhim kho transcriptional gene silencing [49], thiab lub hauv paus txheej txheem kho Pelota tiv thaiv geminiviruses tsis paub.
Cov pov thawj ntxiv tias geminiviruses qhib ETI yog muab los ntawm kev tshawb fawb ntawm geminivirus inducers thiab suppressors ntawm HR thiab ETI-zoo li cov lus teb (Daim duab 3). Geminivirus kab mob induces hypersensitive teb (HR) thiab senescence-txog noob yam tsis tsim ib tug pom cell tuag phenotype [50]. Tshwj xeeb tshaj yog, geminivirus proteins Rep, NSP, thiab V2 tau pom tias ua rau HR [51–53].
Hauv qhov sib piv, qee cov geminiviral proteins muaj lub peev xwm ntawm HR-zoo li cell tuag [19]. C2 protein los ntawm papaya leaf curl virus (PaLCuV) thiab paj rwb nplooj curl Kokhran virus (CLCuKoV) tau pom tias inhibit V2-mediated HR [53]. Ib yam li ntawd, TYLCV kab mob alleviates cell tuag nyob rau hauv txiv lws suav nroj tsuag, inactivation ntawm tshav kub shock protein 90 (HSP90) thiab suppressor ntawm G2 allele ntawm Skp1 (SGT1) [54].
Tseem, TYLCV-mediated cell tuag suppression mechanism tsis paub. Nyob rau hauv sib piv, lub hauv paus txheej txheem rau lub cell tuag-suppressing kev ua ntawm C4 los ntawm lws suav nplooj curl Yunnan tus kab mob (TLCYnV) twb tsis ntev los no uncovered [55]. C4 cuam tshuam nrog HIR1, cuam tshuam HIR1 tus kheej-oligomerization, thiab txhawb nqa nws cov degradation, yog li inhibiting HIR1-koom haum HR thiab nce tus kab mob pathogenicity. Txawm hais tias ob peb kab ntawm cov pov thawj qhia tau hais tias ob qho tib si monopartite thiab bipartite begomoviruses induce thiab suppress HR, cov txheej txheem ntawm ETI ua kom thiab tsub kom los ntawm geminiviruses tseem nyob deb ntawm kev nkag siab.
4. NIK1-Mediated Antiviral Signaling thiab Crosstalk nrog Antibacterial Immunity
NIK1 tau pom thawj zaug hauv cov txiv lws suav ua lub hom phiaj ntawm NSP los ntawm tus kab mob txiv lws suav golden mosaic (TGMV). Nws transduces ib tug antiviral teeb liab uas culminates nyob rau hauv lub suppression ntawm lub ntiaj teb no host translation [16,18]. NIK1 kev sib cuam tshuam nrog NSP yog khaws cia ntawm NIKs los ntawm ntau lub tswv yim, xws li Arabidopsis, taum pauv, thiab txiv lws suav, thiab NSPs los ntawm cov kab mob begomoviruses sib txawv, suav nrog cov kab mob cab nplooj curl (CabLCV), TGMV, txiv lws suav crinkle daj nplooj kab mob (TCrYLV), thiab txiv lws suav daj. Spot Virus (ToYSV) [30,56]. Tsis ntev los no, NIK2 los ntawm Arabidopsis tau pom tias ua haujlwm raws li NIK1 paralog hauv cov kab mob tiv thaiv kab mob [7,27].
Txawm hais tias NIK1-kev kho mob tshuaj tiv thaiv kab mob tau raug suav tias yog ib qho kev qhia tshiab rau cov nroj tsuag tiv thaiv kab mob, nws nthuav tawm ntau yam PTI [57]. Ua ntej, NIK1 yog cov qauv tsim muaj feem xyuam nrog cov PTI coreceptors BAK1, SERK4, thiab SERK1, uas tso saib cov npe kinase tau txais txiaj ntsig zoo thiab kev teeb tsa zoo sib xws ntawm LRR extracellular domain [27] (Daim duab 2).
Thib ob, NIK1 kev ua kom muaj zog yuav tsum muaj phosphorylation ntawm threonine-474, ib txoj haujlwm txuag ntawm cov chaw ua haujlwm ntawm BAK1, SERK4, thiab SERK1, uas tej zaum yuav qhia tau tias muaj ib qho zoo sib xws rau kinase activation [58]. Thib peb, NIK1 yog qhib los ntawm begomovirusderived nucleic acids, zoo li PTI ua kom lub cev los ntawm kab mob PAMPs [7,22]. Thaum kawg, NIK1-kev kho cov kab mob sib kis tau cuam tshuam los ntawm begomovirus-encoded NSP, ua kom tiav cov ntsiab lus tias kev ua tiav ETI yuav yog nyob ntawm PTI kev tawm tsam los ntawm cov kab mob kis [7,11]. Piv txwv ntawm PTI suppressors los ntawm lwm cov kab mob nroj tsuag muaj xws li CP los ntawm plum pox virus (PPV), Potyvirus genus [8], P6 los ntawm cauliflower mosaic virus (CaMV), Caulimovirus genus [59], thiab MP los ntawm dib mosaic virus (CMV), Cucumovirus genus [9]. Yog li ntawd, zoo ib yam li cov kab mob classic PTI, NIK1-kev kho cov kab mob sib kis tau qhib thiab ua rau muaj kev cuam tshuam los ntawm tus kab mob begomovirus.
Txawm hais tias qhov zoo sib xws ntawm PTI thiab NIK1 signaling, cov xwm txheej hauv qab ntawm receptor / coreceptor activation thiab kev sib dhos ntawm cov lus teb tiv thaiv txawv heev. Feem ntau, PTI kev ua kom ua rau ROS tsub zuj zuj raws li cov lus teb ntxov, tom qab ntawd los ntawm kev ua haujlwm ntawm MAP Kinases, induction ntawm PTI-associated tiv thaiv noob, ethylene thiab salicylic acid synthesis, thiab callose deposition [5]. Sib txawv ntawm PTI cov lus teb tiv thaiv kab mob, NIK1- nruab nrab cov tshuaj tiv thaiv kab mob ua rau muaj kev cuam tshuam ntawm tus tswv tsev neeg txhais lus thoob ntiaj teb (Daim duab 4) [60].
Raws li tus qauv siv tshuab rau NIK1 kev ua kom muaj zog, thaum kis kab mob, begomovirus-derived nucleic acids ua haujlwm raws li cov kab mob PAMPs uas ua rau NIK1 dimerization nrog nws tus kheej, NIK2, lossis ib qho tsis paub receptor (Daim duab 4) [7,30]. Raws li cov tswv cuab ntawm cov tsev neeg II ntawm LRRII-RLK nrog cov kev teeb tsa ectodomain zoo ib yam li PTI coreceptors, Arabidopsis NIK1 thiab nws cov paralog NIK2 tuaj yeem ua haujlwm ua cov coreceptors ntawm cov tsis paub nucleic acid-sensing PRRs [6]. Ib qho kev ua haujlwm ntawm lub cev tiv thaiv kab mob ua rau NIK1 phosphorylation ntawm Thr-474, qhov tseem ceeb rau NIK1 kinase activation los pib teeb liab [41,58]. Activated NIK1 mediates lub phosphorylation ntawm downstream tivthaiv ribosomal protein (RP) L10, uas nyob rau hauv lem txhawb nws translocation mus rau lub nucleus [61,62]. NIK1 tuaj yeem tswj tau qhov twg ntawm RPL10 phosphorylation los ntawm kev sib txuas autophosphorylation ntawm qhov ua haujlwm antagonistic Thr-469 residue [41].
Nyob rau hauv lub nucleus, phosphorylated RPL10 cuam tshuam nrog cov transcriptional repressor L10- interacting Myb-domain-muaj protein (LIMYB) kom tag nrho repress cov kev qhia ntawm translational machinery-related noob, ua rau suppression ntawm lub ntiaj teb no protein synthesis [18]. Viral mRNAs tsis tuaj yeem khiav tawm ntawm tus tswv tsev txhais lus tsuj; lawv tsis txhais tau zoo, cuam tshuam tus kab mob [16,18]. Kev tiv thaiv kev ua kom muaj zog ntawm txoj kev tiv thaiv, NSP cuam tshuam nrog kinase domain los tiv thaiv NIK1 phosphorylation thiab ua kom muaj zog, tsim kom muaj ib puag ncig zoo rau kev kis kab mob begomovirus [30,41,61]. Yog li ntawd, NIK1-kev kho mob tshuaj tiv thaiv kab mob yog ib qho evolutionarily suppressed host tiv thaiv los ntawm begomoviruses.
Daim duab 4. NIK1-kev kho mob antiviral signaling thiab crosstalk nrog tshuaj tiv thaiv kab mob: Hauv cov hlwb uas tsis muaj kab mob, (1) NIK1 pas dej ua ke yog khi rau PRR FLS2 thiab PTI coreceptor BAK1 los tiv thaiv autoimmunity. (2) Thaum pib kis tus kab mob, CP-bound ssDNA raug thauj mus rau lub nucleus rau kev rov ua dua ntawm tus kab mob genome thiab kev hloov pauv ntawm cov noob kab mob. (3)
Nyob rau hauv cov kab mob, geminivirus-derived nucleic acids ua raws li tus kab mob PAMPs thiab induce lub dimerization ntawm NIK1 nrog nws tus kheej los yog nrog lwm yam tsis paub receptor (PRR-zoo li) rau transphosphorylation ntawm kinase intracellular domains. NIK1 phosphorylation ntawm Thr-474 activates kinase uas nyob rau hauv lem mediates RPL10 phosphorylation. (4) Phosphorylated RPL10 yog redirected rau lub nucleus, qhov twg nws cuam tshuam nrog LIMYB los tawm tsam qhov kev qhia ntawm ribosomal proteins (RBs) thiab translational machinery-related genes, culminating in suppressing host and viral mRNA translation. (5) NSP tiv thaiv qhov kev tiv thaiv kab mob no los ntawm kev khi rau NIK1 kinase domain, yog li tiv thaiv NIK1 phosphorylation thiab ua kom muaj zog. (6) Cov kab mob kab mob (Pseudomonas spp) muab cov kab mob apoplastic PAMP flagellin (flg22), uas tau lees paub los ntawm PRR FLS2, yog li inducing tsim cov active tiv thaiv complex FLS2-BAK1 los pib PTI. FLS2-BAK1- khi NIK1 yog tom qab ntawd phosphorylated ntawm Thr-474 los ntawm kev qhib BAK1, (7) txhim kho NIK1 kev tawm tsam ntawm PTI tab sis (8) ua kom NIK1- kho kom haum xeeb antiviral signaling uas ua rau muaj kev tawm tsam ntawm kev txhais lus thoob ntiaj teb. (9) C4 los ntawm geminiviruses khi rau FLS2 thiab inhibits PTI cov lus teb thaum ntxov thiab tom qab NIK1 ua kom tsim tau ib puag ncig uas nyiam cov kab mob. Cov lus nug cov qhab nia qhia txog qhov xwm txheej tsis tau qhia meej lossis tsis paub. Daim duab tau tsim nrog BioRender 101.
NIK1-kev tiv thaiv kab mob sib kis sib txuas nrog PTI vim NIK1 thiab NIK2 kuj ua haujlwm raws li PTI suppressors (Daim duab 4) [17]. Inactivation ntawm NIK1 lossis NIK2 hauv Arabidopsis txhim kho PTI kev ua kom muaj zog thiab ua rau muaj kev tiv thaiv rau Pseudomonas syringae, ib qho kev sib txawv phenotype uas pom los ntawm NIK-overexpressing kab. Raws li ib txwm muaj, NIK1 cuam tshuam nrog flagellin receptor FLS2 thiab nws cov coreceptor BAK1 cuam tshuam nrog kev tiv thaiv kab mob thiab tiv thaiv kev tiv thaiv autoimmunity (Daim duab 4).
Cov kab mob PAMP flagellin, los yog nws cov nquag peptide flg22, cuam tshuam nrog FLS2 thiab induces dimerization nrog BAK1, ua rau tsim ib qho kev tiv thaiv kab mob kom pib PTI signaling. FLS2-kev ua kom sib haum xeeb ntawm BAK1 txhawb nqa NIK1 phosphorylation ntawm qhov chaw ua kom Thr-474, uas txhawb nqa ntxiv NIK1 kev sib cuam tshuam nrog FLS2 thiab BAK1 kom tswj tau qhov kev ua kom PTI thiab tig ua kom lub NIK 1-kev kho mob tshuaj tiv thaiv kab mob. Flg22-induced NIK1 phosphorylation yuav tsum muaj ob qho tib si PRR FLS2 thiab nws cov coreceptor BAK1, qhia tias NIK1 ua rau qis qis ntawm receptor signaling. Cov txiaj ntsig no tau lees paub qhov kev sib cav tias NIK1 tuaj yeem ua haujlwm ua tus coreceptor hauv ligand-induced activation ntawm ib qho tseem tsis tau paub txog nucleic acid-sensing receptor (PRR) txhawm rau txhawm rau NIK1-kev tiv thaiv kab mob sib kis. Lawv kuj qhia tias cov kab mob PAMP tuaj yeem ua rau muaj kab mob begomovirus los ntawm NIK1 phosphorylation, uas tuaj yeem txo NSP inhibition. Ntau cov ntaub ntawv pov thawj qhia tau hais tias phosphorylation-mediated activation ntawm NIK1 yog downstream ntawm NIK1 inhibition los ntawm begomovirus-encoded NSP.
Ua ntej, NSP-binding site ntawm NIK1 maps nyob rau hauv ib qho 80 amino acid stretch overlapping lub ua kom lub voj thiab lub kinase activation site (Thr474) [30]. Thib ob, kev hloov pauv ntawm Thr-474 qhov chaw ua haujlwm nrog ib qho aspartate tsim ib qho phosphomimetic mutant uas tsis yog inhibited los ntawm NSP [41]. Thaum kawg, overexpression ntawm phosphomimetic mutant NIK1-T474D nyob rau hauv Arabidopsis thiab txiv lws suav nroj tsuag muaj ntau yam tiv thaiv kab mob begomovirus [16,18]. Yog li ntawd, kev kho tus tswv tsev nrog cov kab mob PAMP flg22 ua ntej tus kab mob begomovirus tuaj yeem ua rau muaj txiaj ntsig zoo NIK1- cov tshuaj tiv thaiv kab mob sib kis los ntawm kev hla NSP inhibition. Qhov kev sib cuam tshuam ntawm kev tiv thaiv kab mob no tso cai rau cov kab mob ua kom muaj kev tiv thaiv kab mob tiv thaiv kab mob, yog li tiv thaiv ntau yam kab mob los ntawm cov kab mob los ntawm ntau lub nceeg vaj xav tias kom tsis txhob muaj kev sib tw.
5. Cov lus xaus
Cov kev tshawb fawb tsis ntev los no tau pom tias cov nroj tsuag siv ob theem ntawm kev tiv thaiv kab mob hauv lub cev, PTI thiab ETI, txhawm rau tiv thaiv kab mob, thiab geminiviruses tsis yog qhov zam. Txawm li cas los xij, pov thawj rau geminiviral PTI thiab ETI tso siab rau cov kev soj ntsuam uas tus kab mob geminivirus tuaj yeem ua rau thiab ua rau PTI thiab ETI zoo li cov lus teb. Tsis tas li ntawd, qee qhov geminiviral proteins ua haujlwm li PTI suppressors, thiab tsawg kawg yog ib qho kev sib cais ntawm cov noob rau TYLCV qhia txog cov txheej txheem NL-LRR classical ntawm intracellular R proteins uas elicits ETI. Txawm li cas los xij, peb txoj kev paub txog geminiviral PTI thiab ETI cov txheej txheem tseem yog qhov tseem ceeb, thiab ob peb kauj ruam hauv PTI thiab ETI ua kom thiab teb tsis paub. Piv txwv li, peb tsis paub tus kheej ntawm geminiviral PAMPs uas yuav ntxias PTI, thiab cov cognate PRRs tsis tau txheeb xyuas.
Tsis tas li ntawd, lub repertoire ntawm PTI geminiviral suppressors yog txwv rau C4 thiab tej zaum mus rau C1 thiab NSP. NSP-suppressed NIK1-kev kho mob tshuaj tiv thaiv kab mob tau pom tias muaj kev sib txuas lus nrog cov kab mob PTI. Ntxiv nrog rau kev txwv tsis pub neeg txhais lus thoob ntiaj teb, NIK1 kev ua kom ua rau muaj kev cuam tshuam ntawm cov kab mob PTI, uas txwv tsis pub ntxiv kev siv NIK txuas ntxiv 1- kho cov kab mob tiv thaiv kab mob los tswj tus kab mob. Kev rho tawm ntawm geminiviral PAMPs thiab cognate PRRs, geminiviral effectors (Avr genes), thiab NL-LRR lub hom phiaj yuav pab kom paub meej lub tswv yim ntawm ETI ua kom thiab PTI kev tawm tsam ua lub hauv paus rau kev nkag siab txog kev hloov pauv siab ua haujlwm ntawm geminivirus thiab kev sib cuam tshuam ntawm tus tswv tsev. Qhov kev nkag siab no thaum kawg yuav pab txiav txim siab yuav ua li cas siv lub cev tiv thaiv kab mob los tswj tus kab mob geminivirus.
Tus sau kev koom tes:
MAF thiab RMT tau sau thawj tsab ntawv ntawm cov ntawv sau. EPBF tau tsim thiab saib xyuas cov ncauj lus tshuaj xyuas. Txhua tus kws sau ntawv tau pab txhawb rau tsab xov xwm. Txhua tus kws sau ntawv tau nyeem thiab pom zoo rau cov ntawv luam tawm ntawm cov ntawv sau.
Nyiaj txiag:
Txoj haujlwm no tau txais nyiaj ib nrab los ntawm CAPES nyiaj txiag code 001, CNPq, FAPEMIG, thiab National Institute of Science thiab Technology hauv kev sib cuam tshuam ntawm tsob nroj-Pest. MA yog tus tau txais kev pabcuam CAPES kawm tiav thiab RMT tau txais kev txhawb nqa los ntawm FAPMIG kawm tiav kev sib raug zoo.
Kev tsis sib haum xeeb ntawm kev txaus siab:
Cov kws sau ntawv tshaj tawm tsis muaj kev sib cav txog kev txaus siab.
Cov ntaub ntawv
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Marco Aurélio Ferreira, Ruan M. Teixeira thiab Elizabeth PB Fontes.
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