Lub luag haujlwm tseem ceeb ntawm Motile Epidermal Chloroplasts hauv Kev Tiv Thaiv Kab Mob
May 19, 2023
Abstract:
Cov nroj tsuag epidermis muaj atypical me me chloroplasts. Txawm li cas los xij, lub luag haujlwm ntawm lub cev ntawm cov organelle no tsis meej meej piv rau cov mesophyll chloroplasts loj, cov haujlwm paub zoo uas yog photosynthesis. Txawm hais tias kev paub txog kev koom tes ntawm chloroplasts hauv cov nroj tsuag tiv thaiv kab mob tau nthuav dav rau hnub no, qhov sib txawv ntawm cov kab mob epidermal thiab mesophyll chloroplasts yog dhau ntawm qhov kev tshawb fawb no. Muab lub luag haujlwm ntawm cov nroj tsuag epidermis ua ib qho kev cuam tshuam rau ib puag ncig kev ntxhov siab, nrog rau cov kab mob tawm tsam, thiab kev tiv thaiv kev tiv thaiv kev ua haujlwm ntawm chloroplasts, cog kev tiv thaiv kev tshawb fawb ntawm epidermal chloroplasts yog qhov tshwm sim. Cov kev tshawb fawb tsis ntev los no tau qhia txog kev txav mus los ntawm epidermal chloroplasts teb rau cov kab mob fungal thiab oomycete. Tsis tas li ntawd, epidermal chloroplast-txuas nrog cov proteins thiab cov xwm txheej ntawm tes uas txuas nrog cov kab mob epidermal tiv thaiv kab mob. Hauv qhov kev tshuaj xyuas no, kuv tau tsom mus rau qhov kev nce qib tsis ntev los no hauv epidermal chloroplast-mediated cog kev tiv thaiv kab mob.
Ntsiab lus: epidermal chloroplast; chloroplast zog; stromule; nroj tsuag kab mob; nkag mus; nonhost kuj; preinvasive tiv thaiv; Arabidopsis thaliana; Nicotiana benthamiana; CHAW 1.
Kev tiv thaiv cog qoob loo thiab kev tiv thaiv tib neeg yog ob txoj kev tiv thaiv sib txawv, thiab muaj qhov sib txawv pom tseeb:
1. Cov kab mob sib txawv los tawm tsam: cov nroj tsuag feem ntau tawm tsam cov kab mob lom xws li kab mob, fungi, thiab kab mob, thaum tib neeg kev tiv thaiv tsuas yog tiv thaiv kab mob thiab kab mob.
2. Cov txheej txheem tiv thaiv kab mob sib txawv: cog kev tiv thaiv tsuas yog tiv thaiv kab mob los ntawm kev sib cais ntawm lub cev, tshuaj tiv thaiv kab mob, thiab kev tswj cov noob, thaum tib neeg kev tiv thaiv tsuas yog tiv thaiv kab mob los ntawm macrophages, T hlwb, thiab B hlwb.
3. Kev tiv thaiv kab mob sib txawv: Kev tiv thaiv cog qoob loo siv tib txoj kev tiv thaiv tiv thaiv txhua tus kab mob, uas yog, tsis muaj kev tiv thaiv tshwj xeeb, thaum tib neeg kev tiv thaiv tshwj xeeb, muaj peev xwm lees paub thiab tsim cov tshuaj tiv thaiv kab mob tiv thaiv kab mob tshwj xeeb.
4. Cov kab mob tiv thaiv kab mob sib txawv: Cov nroj tsuag tsis muaj peev xwm ntawm kev tiv thaiv kab mob, uas yog, thaum lawv tau tawm tsam qee yam kab mob, lawv tseem yuav tsum rov pib lub cev tiv thaiv kab mob rau lwm zaus lawv ntsib tib yam kab mob. Tib neeg lub cev tiv thaiv kab mob muaj peev xwm ntawm lub cev tiv thaiv kab mob, uas yog, tom qab tiv thaiv kab mob, nws yuav muaj lub cev tiv thaiv kab mob ntev ntev rau cov kab mob no, thiab lwm zaus nws ntsib cov kab mob qub dua, nws tuaj yeem ua kom lub cev tiv thaiv kab mob sai sai. , yog li zoo tswj cov kab mob kis. Yog li nyob rau hauv kev sib piv, peb tib neeg kev tiv thaiv kuj tseem ceeb heev. Peb yuav tsum tsom ntsoov rau kev txhim kho peb txoj kev tiv thaiv. Cistanche deserticola tuaj yeem ua tau. Cov polysaccharides hauv cov nqaij tuaj yeem tswj hwm kev tiv thaiv kab mob ntawm tib neeg lub cev tiv thaiv kab mob thiab txhim kho kev ntxhov siab ntawm lub cev tiv thaiv kab mob. , Txhim kho cov kab mob bactericidal ntawm lub cev tiv thaiv kab mob.
Nyem cistanche deserticola ntxiv
1. Taw qhia
Lub epidermis ntawm ntau cov kab mob, nrog rau cov nroj tsuag, ua ib qho kev tiv thaiv los tiv thaiv ntau yam kev ntxhov siab xws li kev hloov hauv ib puag ncig sab nraud thiab cov kab mob tawm tsam. Los ntawm qhov kev xav ntawm cov kab mob, kev tawg los ntawm cov kab mob epidermal ntawm cov nroj tsuag yog qhov tseem ceeb thawj kauj ruam hauv kev cuam tshuam; tshwj xeeb, cov kab mob fungal thiab oomycete yuav tsum nkag mus ncaj qha rau hauv cov nroj tsuag epidermis thiab tsim cov kab mob hyphae sab hauv kom muaj kev vam meej. Piv txwv li, anthracnose fungi Colletotrichum hom thiab nplej moj tej tawg fungus Pyricularia oryzae (syn. Magnaporthe oryzae) tsim ib tug melanized dome-zoo li tus cell hu ua appressorium rau ntawm cov nroj tsuag nto los tsim ib tug penetration peg nyob rau hauv lub epidermis ntawm tus tswv nroj tsuag, ua raws li invasive. hyphal extension thiab qhov tshwm sim ntawm tus kab mob puas tsuaj [1].
Txawm li cas los xij, yog tias cov nroj tsuag tsis yog tus tswv, cov kab mob fungal no tuaj yeem tsim melanized appressorium, tab sis tsis tuaj yeem nkag mus rau hauv cov kab mob epidermal vim yog cov kab mob ua ntej tsis muaj zog tiv thaiv (NHR) ntawm cov nroj tsuag, uas feem ntau muab kev ruaj khov, muaj zog, thiab broadspectrum kev tiv thaiv thiab tiv thaiv zoo. kev ntxeem tau ntawm ntau cov kab mob uas tsis muaj kab mob thiab oomycetes hauv kev sib cuam tshuam tsis sib xws [2,3]. Nyob rau hauv tus qauv brassicaceous cog Arabidopsis thaliana, nws tau raug tshaj tawm tias ntau txoj hauv kev tiv thaiv kab mob thiab cov khoom siv los ntawm kev xa tawm ntawm epidermal NHR nyob rau hauv ntau yam tawm tsam kev nkag mus ntawm nonadapted fungal thiab oomycete pathogens, xws li Blumeria graminis f. sp. hordei [4–12], Colletotrichum tropicale [13–16], P. oryzae [17–22], thiab Phytophthora infestans [6,7]. Txawm li cas los xij, kev nkag siab tiav ntawm lub hauv paus molecular ntawm preinvasive NHR tseem tsis pom. Txhawm rau kom tau txais kev nkag siab zoo ntawm NHR, nws yog ib qho tseem ceeb los txheeb xyuas cov xwm txheej tsis paub txog kev tiv thaiv kab mob thiab cov khoom hauv cov nroj tsuag epidermis.
Hauv cov nroj tsuag siab dua, tshwj tsis yog qee cov nroj tsuag, xws li haus luam yeeb, nws tau ntseeg ntev tias tsis muaj chloroplasts muaj nyob hauv cov kab mob epidermal uas tsis yog cov neeg saib xyuas [23–26]. Feem ntau cov kev tshawb fawb ntawm chloroplast tsom rau cov hlwb mesophyll, qhov twg ntau cov chloroplasts loj heev sib txawv heev rau photosynthesis [27–29]. Txawm li cas los xij, chlorophyll-muaj atypical me me chloroplasts kuj tau pom nyob rau hauv cov epidermal pavement hlwb ntawm A. thaliana, txawm hais tias thylakoids, uas yog lub luag hauj lwm rau lub teeb-dependent photosynthesis tshwm sim, tsis zoo tsim [30–36]. Yog li ntawd, lub luag haujlwm ntawm lub cev ntawm epidermal chloroplasts yog lub ntsiab lus tseem ceeb hauv kev cog qoob loo.
Chloroplast paub tias ua haujlwm li ib puag ncig sensors tiv thaiv ntau yam kab mob biotic thiab abiotic stresses [37,38]. Piv txwv li, rau cov nroj tsuag tiv thaiv kab mob tiv thaiv kab mob, cov tub txib theem nrab, xws li reactive oxygen hom (ROS) thiab calcium (Ca2 ntxiv), thiab cov precursors ntawm phytohormones, xws li salicylic acid (SA), jasmonic acid, thiab abscisic acid, yog Tag nrho cov muab tau los ntawm chloroplasts [39–42]. Muaj ntau qhov kev tshuaj xyuas zoo txog lub luag haujlwm ntawm chloroplasts hauv cov nroj tsuag tiv thaiv kab mob. Kuv xa cov neeg nyeem mus rau cov kev tshuaj xyuas tsis ntev los no, uas yuav muab cov ntsiab lus ntawm chloroplast hais txog cov lus teb tiv thaiv nroj tsuag [43–47].
Txawm li cas los xij, muaj qee qhov kev tshawb fawb txog kev tiv thaiv cov nroj tsuag uas tsom mus rau epidermal chloroplasts, thiab kev ua haujlwm sib txawv ntawm epidermal thiab mesophyll chloroplasts hauv cov nroj tsuag tiv thaiv kab mob tsis tau txais kev saib xyuas ntau. Tsis ntev los no, qee cov ntawv tshaj tawm tau qhia txog kev koom tes ntawm epidermal chloroplasts hauv cov nroj tsuag tiv thaiv kab mob. Hauv qhov kev tshuaj xyuas no, lub hom phiaj tseem ceeb yog ntawm cov tshuaj chloroplasts epidermal, thiab cov kev kawm tsis ntev los no hauv cov nroj tsuag epidermal tiv thaiv kab mob tau piav qhia.
2. Epidermal Chloroplast Response Tswj kev nkag mus ntawm cov kab mob fungal hauv A. thaliana
2.1. Intracellular Movements ntawm Epidermal Chloroplast nyob rau hauv Teb rau Fungal Pathogens hauv A. thaliana
Nyob rau hauv tsis tu ncua ntawm A. thaliana, epidermal chloroplasts tsis pom nyob ze ntawm lub Upper periclinal phab ntsa (nto) ntawm pavement hlwb, vim hais tias lawv feem ntau yog positioned nyob rau hauv qis periclinal (hauv qab) thiab anticlinal phab ntsa. Hauv tsab ntawv tshaj tawm tsis ntev los no, kuv pom tias epidermal chloroplasts tau tshwm sim nyob rau saum npoo ntawm pavement hlwb teb rau Colletotrichum fungi nrog melanized appressorium tsim, thiab hu ua qhov tshwm sim no epidermal chloroplast teb (ECR) (Daim duab 1) [48].
Interestingly, ECR occurs more strongly against nonadapted fungi such as the Japanese flowering cherry pathogen Colletotrichum nymphaea PL1-1-b, the cosmos pathogen Colletotrichum fioriniae CC1, and the apple pathogen Colletotrichum siamense MAF1, compared to the adapted Brassicaceae pathogen Colletotrichum higginsianum Abr1-5, which readily penetrates and infects wild-type plants of A. thaliana [48]. Furthermore, the frequency of ECR varies according to the nonadapted fungal strains; the order of ECR-inducing ability in the wild-type plant is as follows: C. nymphaea PL1-1-B > C. fioriniae CC1 > C. siamense MAF1, while two other nonadapted strains, hau tree pathogen C. siamense COC4, and cucurbit pathogen Colletotrichum orbiculare 104-T, do not trigger ECR in the wild-type plant [48]. Importantly, C. siamense COC4 and C. orbiculare 104-T sufficiently induce ECR in plants with a mutation in the PEN2 gene [48], which encodes an atypical myrosinase that works as a core preinvasive NHR contributor against many fungal and oomycete pathogens [6,9,13,17,48–50], although these two nonadapted pathogens cannot invade pen2 mutants. The frequencies of ECR in the pen2 mutant against C. fioriniae CC1 and C. siamense MAF1 are also higher than those in the wild-type plant, whereas other single mutants of immune-related genes such as EDR1, GSH1 EDS5, and CAS have little effect on ECR [48]. These results suggest that ECR is preferentially activated when PEN2-based antifungal preinvasive defense becomes ineffective in epidermal cells. Similarly, in penetration tests on multiple Arabidopsis mutants with many immune-related mutations, these five nonadapted Colletotrichum strains displayed the differential abilities to overcome preinvasive NHR in the same order as the ECR-inducing ability; C. nymphaea PL1-1-B and C. fioriniae CC1 can invade the epidermis of the wild-type plant to some degree and more of the pen2 mutant, whereas C. siamense MAF1, COC4, and C. orbiculare 104-T can break through the epidermal NHR only in the presence of pen2 mutation or with additional mutations such as edr1, gsh1, eds5, and cas (Figure 2, Redrawn from [51]) [48,51]. Thus, there is a tight link between ECR induction in plants and the invasion ability of the nonadapted Colletotrichum fungi, which implies the involvement of ECR in epidermal preinvasive NHR, because these fungal pathogens are incompatible with and definitely cannot infect A. thaliana. The contribution of ECR to preinvasive NHR is described in a later section. Int. J. Mol. Sci. 2022, 23, x FOR PEER REVIEW 3 of 17 readily penetrates and infects wild-type plants of A. thaliana [48]. Furthermore, the frequency of ECR varies according to the nonadapted fungal strains; the order of ECR-inducing ability in the wild-type plant is as follows: C. nymphaea PL1-1-B > C. fioriniae CC1 >C. siamense MAF1, thaum ob hom kab mob uas tsis hloov pauv, muaj tsob ntoo kab mob C. siamense COC4, thiab cucurbit pathogen Colletotrichum orbiculare 104-T, tsis txhob ua rau ECR hauv cov nroj tsuag qus [48].
Tseem ceeb, C. siamense COC4 thiab C. orbiculare 104-T txaus induce ECR nyob rau hauv cov nroj tsuag nrog ib tug hloov nyob rau hauv lub PEN2 noob [48], uas encodes ib tug atypical myrosinase uas ua hauj lwm raws li ib tug core preinvasive NHR contributor tiv thaiv ntau fungal thiab oomycete. Cov kab mob [6,9,13,17,48–50], txawm hais tias ob hom kab mob no tsis tuaj yeem nkag mus rau pen2 mutants. Cov zaus ntawm ECR nyob rau hauv pen2 mutant tiv thaiv C. fioriniae CC1 thiab C. siamense MAF1 kuj siab dua cov nyob rau hauv cov qus-hom nroj tsuag, whereas lwm yam mutants ntawm kev tiv thaiv kab mob xws li EDR1, GSH1 EDS5, thiab CAS muaj tsawg. cuam tshuam rau ECR [48]. Cov txiaj ntsig no qhia tias ECR yog qhov zoo tshaj plaws qhib thaum PEN2-raws li kev tiv thaiv kab mob tiv thaiv kab mob ua ntej ua tsis tau zoo hauv cov hlwb epidermal.
Ib yam li ntawd, hauv kev ntsuam xyuas nkag mus rau ntau yam Arabidopsis mutants nrog ntau yam kev hloov pauv ntawm kev tiv thaiv kab mob, cov tsib cov kab mob Colletotrichum nonadapted tau nthuav tawm lub peev xwm sib txawv los kov yeej preinvasive NHR nyob rau hauv tib txoj kev raws li ECR-inducing peev xwm; C. nymphaea PL1-1-B thiab C. fioriniae CC1 tuaj yeem cuam tshuam cov epidermis ntawm cov nroj tsuag qus mus rau qee qhov kev kawm thiab ntau dua ntawm pen2 mutant, whereas C. siamense MAF1, COC4, thiab C. orbiculare {{8 }}T tuaj yeem cuam tshuam los ntawm epidermal NHR nkaus xwb nyob rau hauv lub xub ntiag ntawm kev hloov pauv pen2 lossis nrog kev hloov pauv ntxiv xws li edr1, gsh1, eds5, thiab cas (Daim duab 2, Redrawn los ntawm [51]) [48,51]. Yog li, muaj kev sib txuas ntawm ECR induction nyob rau hauv cov nroj tsuag thiab invasion peev xwm ntawm nonadapted Colletotrichum fungi, uas qhia txog kev koom tes ntawm ECR nyob rau hauv epidermal preinvasive NHR, vim hais tias cov fungal pathogens incompatible nrog thiab yeej tsis muaj peev xwm kis tau A. thaliana. Kev koom tes ntawm ECR rau preinvasive NHR yog nonhost Arabidopsis mutants tau raug soj ntsuam raws li melanized appressorium-mediated entry (MAE) cov nqi thiab cais. MAE tus nqi (feem pua) tau suav nrog cov lej hauv qab no: (tus naj npawb ntawm melanized appressoria nrog rau kev tsim ntawm invasive hypha) / (tus naj npawb ntawm melanized appressoria). Adapted C. higginsianum Abr1-5 (MAFF305635) yog qhia raws li kev tswj. Qhov feem pua ntawm cov kab mob fungal nkag tau raws li "−", "ntxiv /-", "ntxiv", "ntxiv ntxiv", "ntxiv ntxiv", "ntxiv ntxiv ntxiv", thiab "ntxiv ntxiv ntxiv ntxiv" yog 0-2 feem pua, 2-10 feem pua, 10-20 feem pua, 20-35 feem pua, 35-60 feem pua, thiab 60-100 feem pua, feem. nd: tsis tau txiav txim. Yuav ua li cas kho los ntawm [51]
ECR tsis yog tshwj xeeb rau Colletotrichum. Epidermal chloroplasts kuj tshwm sim ntawm qhov chaw tom qab inoculation nrog cov nonadapted pathogen P. oryzae, uas qhia tau hais tias melanized appressorium-mediated nkag (MAE)-hom nroj tsuag ntxeem tau tab sis yog phylogenetically distantly cuam tshuam rau Colletotrichum [48]. Qhov zaus ntawm P. oryzae-induced ECR kuj nce hauv tus cwj mem2 mutant. Yog li, ECR tej zaum yuav yog qhov dav-spectrum epidermal teb rau ntau yam kab mob fungal. Lub tswv yim no yog ua raws li qhov kev tshawb pom tshiab uas cov nonadapted fungus Alternaria alternata, uas tsis yog MAE-hom kab mob thiab ua rau melanized conidia, kuj induces ECR nyob rau hauv lub nonhost A. thaliana, tshwj xeeb tshaj yog nyob rau hauv lub xub ntiag ntawm pen2 kev hloov (Daim duab 3) .
2.2. Qhov tshwm sim ntawm Epidermal Chloroplast Teb
Txawm hais tias yuav ua li cas cov nroj tsuag epidermis paub txog cov kab mob fungal thiab induces ECR tseem yog lo lus nug qhib, qee cov ntaub ntawv qhia txog kev sib raug zoo ntawm cov kab mob fungal nkag mus sim cov yam ntxwv thiab ECR induction [48]. Qee qhov kev tshawb fawb tau zoo ntawm cov kab mob uas tau los ntawm cov kab mob yog cov kab mob / cov kab mob sib txuas nrog cov kab mob (PAMPs / MAMPs), uas tau txais txiaj ntsig zoo ntawm cov kab mob dav dav thiab pab txhawb rau lawv txoj kev vam meej [52]. Kev puas tsuaj-txuas nrog cov qauv molecular (DAMPs), cov khoom siv los ntawm tus tswv tsev xws li Pep1 tso tawm extracellularly thaum muaj kab mob ntxeem tau, kuj yog cov lus qhia rau lub cev tiv thaiv kab mob [53]. PAMPs/DAMPs raug lees paub los ntawm cov nroj tsuag qauv lees paub receptors (PRRs), thiab PAMP / DAMP-tsim tiv thaiv kab mob yog pib los ntawm PRR-associated proteins, xws li RLP 23 thiab RLP30, membrane-embedded receptor-like proteins (RLPs), FLS2 thiab BAK1. , membrane-embedded receptor-like kinases (RLKs), thiab BIK1 thiab PBL1, receptor-like cytoplasmic kinases (RLCKs) [54–61].
Kev tshuaj xyuas tshwj xeeb ntawm PRR signaling hauv kev tiv thaiv cov nroj tsuag tau luam tawm los ntawm ntau tus kws tshawb fawb [62-65]. Intriguingly, ECR tsis yog nyob ntawm BAK1, BIK1, PBL1, Pep1-sensing RLKs, PEPR1, thiab PEPR2; yog li ntawd, PAMP thiab/lossis DAMP cov xwm txheej, tsawg kawg yog cov kev kho kom haum xeeb los ntawm cov RLKs thiab RLCKs, tsis koom nrog kev cuam tshuam ntawm ECR [48]. Tsis tas li ntawd, qhov ntau ntawm ECR nyob rau hauv pen2 mutant cog tawm tsam ntau C. orbicular mutants tsis txaus nyob rau hauv ntau cov kauj ruam ntawm invasion-txog morphogenesis qhia tau hais tias qhov tsim ntawm lub penetration peg, ib rab koob zoo li fungal qauv tawm los ntawm appressorium rau kev vam meej nkag mus rau hauv cov nroj tsuag. epidermis, yog qhov tseem ceeb rau ECR induction [48].
Nroj tsuag kab mob fungi thiab oomycetes zais ib qho arsenal ntawm virulence proteins, hu ua effectors, los tswj cov txheej txheem ntawm tes ntawm tes thiab txhawb kev tiv thaiv kab mob. Ntau qhov kev tshuaj xyuas tau luam tawm ntawm cov nyhuv ntawm cov kab mob nroj tsuag [66–70]. Nyob rau hauv Colletotrichum, fluorescently sau npe effector proteins nyiam nyob rau hauv qab pore ntawm lub appressorium rau secretion los ntawm lub penetration peg [71,72]. Qhov kev tso tawm cov nyhuv no yog nyob ntawm v-SNARE SEC22-kev sib haum xeeb hauv txoj hauv kev hauv lub cev hauv cov kab mob fungus [72] thiab, nthuav, cuam tshuam nrog ECR induction [48]. Cov kev soj ntsuam no qhia txog cov neeg muaj peev xwm hauv qab no raws li qhov tshwm sim ntawm ECR: (i) pathogen-derived molecule (s) secreted los ntawm lub penetration peg thaum lub sij hawm fungal nkag mus sim thiab (ii) cog-derived teeb liab (s) generated nyob rau hauv teb rau qib ntawm fungal kev loj hlob hauv kev sim nkag mus. ECR yog ib qho kev tshwm sim thoob ntiaj teb tawm tsam ntau hom kab mob fungal, suav nrog Colletotrichum hom, P. oryzae, thiab A. alternata (Daim duab 3) [48]. Yog li ntawd, cov txiaj ntsig raug txwv rau qhov nqaim ntawm cov kab mob fungal rau lawv tus tswv tsev tshwj xeeb yuav tsis yog ib qho ntawm ECR; es, cov cell phab ntsa-degrading enzymes los yog cov tub ntxhais kev txuag tau zoo heev ntawm ntau yam kab mob fungi lossis cov cim kev puas tsuaj ntawm cov nroj tsuag tsis cuam tshuam nrog Pep1 yuav ua rau ECR hauv cov nroj tsuag epidermis.
2.3. Tus Regulators ntawm Epidermal Chloroplast Teb thiab Preinvasive Nonhost Resistance ntawm Arabidopsis
Loj mesophyll chloroplasts qhia intracellular txav raws li kev siv ntawm lub teeb kom nyab xeeb thiab ua tau zoo photosynthesis; lawv khiav tawm ntawm lub teeb muaj zog thiab tsiv mus rau cov phab ntsa anticlinal ntawm lub xov tooj ntawm tes los tiv thaiv photodamage (kev zam kev teb) thiab txav mus rau lub teeb tsis muaj zog thiab nyob rau ntawm phab ntsa periclinal txhawm rau ua kom cov photosynthetic efficiency (cov lus teb sib xyaw). Ntau qhov kev tswj hwm cov proteins ntawm qhov kev hloov pauv no, hu ua chloroplast yees duab hloov chaw, tau txheeb xyuas hauv cov hlwb mesophyll [73]. Txawm li cas los xij, muaj ob peb tsab ntawv ceeb toom, thiab me ntsis paub txog cov khoom uas tswj cov kev txav ntawm lub cev ntawm epidermal chloroplasts [74,75]. Hauv cov ntsiab lus no, kuv tsis ntev los no tau pom tias ECR hauv cov kab mob epidermal sib koom cov txheej txheem tswj hwm nrog mesophyll chloroplast yees duab txav chaw [48]. Cov actin-binding protein CHUP1 tsim ib qho chloroplast-actin-raws li lub zog quab yuam thiab yog qhov tseem ceeb rau ob qho tib si tsub zuj zuj thiab zam cov lus teb [76-78], thaum lub auxilin-zoo li J-domain protein JAC1 tswj cov tsos thiab ploj ntawm chloroplast-actin filaments. thiab yuav tsum tau rau cov lus teb tsub zuj zuj [79]. Kev hloov kho caj ces A. thaliana qhia tau hais tias overexpression ntawm CHUP1 proteins ua rau muaj zog tsuj ntawm ECR.
Feem ntau, epidermal chloroplasts tsis tshua pom ntawm thaj chaw saum npoo, txawm tias tom qab inoculation nrog nonadapted fungal pathogens [48]. Txawm li cas los xij, qhov overexpression ntawm JAC1 los yog tshem tawm ntawm CHUP1 ua rau lub constitutive positioning ntawm epidermal chloroplasts nyob rau saum npoo, tsis hais fungal inoculation [48]. Yog li, CHUP1 thiab JAC1 tswj qhov tsis zoo thiab zoo, raws li txoj cai, qhov chaw ntawm epidermal chloroplasts ntawm thaj tsam ntawm cov pavement hlwb (Daim duab 4). Nws yog ib qho tseem ceeb uas inoculation ntawm chup1 mutant nrog siab concentration ntawm nonadapted fungi tsuas yog me ntsis nce cov pej xeem ntawm nto chloroplasts, yog li qhia qhov tsis zoo ntawm ECR los ntawm depletion ntawm CHUP1 protein. Nyob rau hauv sib piv, phototropin 1 thiab 2, uas yog xiav lub teeb receptors lub luag hauj lwm rau chloroplast yees duab txav txav [80-82], yog dispensable rau ua kom ECR [48]. Yog li ntawd, chloroplast daim duab qhov chaw thiab ECR muaj qhov sib txawv stimulus paub tshuab rau lub teeb thiab pathogens, raws li, tab sis qhia tsawg kawg yog ib co downstream kev cai Cheebtsam. Los ntawm qhov kev pom ntawm qhov sib txawv ntawm mesophyll thiab epidermal chloroplasts, versatility ntawm lub adaptive systems tiv thaiv txawv ib puag ncig kev nyuaj siab tej zaum yuav muaj kev cuam tshuam lub elaborate ciaj sia taus lub tswv yim ntawm cov nroj tsuag.
Seb ECR txhawb kev cog kev tiv thaiv kab mob hauv cov kab mob epidermal ntawm A. thaliana yog ib lo lus nug uas ntxim nyiam. Txhawm rau kom paub meej txog kev koom tes ntawm ECR hauv preinvasive NHR, cov teebmeem ntawm ECR kev puas tsuaj los ntawm overexpressing los yog depleting CHUP1 proteins ntawm MAE tus nqi ntawm ntau yam nonadapted fungi nyob rau hauv cov nroj tsuag epidermis tau raug soj ntsuam [48]. C. nymphaea PL1-1-B, uas nthuav tawm qhov tsis sib xws nrog A. thaliana piv rau lwm cov kab mob uas tsis tau hloov pauv, tau pom tias muaj MAE ntau dua hauv cov nroj tsuag uas tsis muaj ECR (Daim duab 2) [48]. Ntxiv mus, hauv cov nroj tsuag nrog cwj mem2 lossis lwm yam kev hloov pauv ntxiv xws li edr1, gsh1, eds5, thiab cas, qhov nce lossis txo qis hauv cov qib CHUP1 proteins txhawb MAE tus nqi ntawm C. fioriniae CC1, C. siamense MAF1, COC4, thiab C. . orbiculare 104-T (Daim duab 2) [48]. Cov txiaj ntsig zoo sib xws tau txais rau MAE tus nqi ntawm P. oryzae. Raws li, ECR yog cov lus teb kev ntxhov siab ntawm cov nroj tsuag, los ntawm qhov preinvasive NHR yog txhim kho nyob rau hauv epidermal hlwb thaum cov kab mob fungal sim ntxeem tau los ntawm kev nkag mus peg.
Raws li tau piav qhia, ECR tau ua kom zoo dua thaum tsis muaj PEN2-txoj kev tiv thaiv. Ib yam li ntawd, qhov inoculation ntawm nonadapted C. fioriniae CC1 kuj induced kev qhia ntawm ntau yam kev tiv thaiv cov noob, xws li PAD3, CYP79B2, CYP71A13, PDF1.2a, MYB51, PR1, FRK1, thiab NHL10, tsuas yog nyob rau hauv cov ntaub ntawv ntawm muv. Txawm hais tias qhov pib ntawm ECR tau qis dua qhov ntawd rau qhov kev qhia ntawm cov noob no [48]. Raws li cov txheej txheem txheej ntawm cov nroj tsuag NHR, ECR yog ib qho ntawm cov lus teb tiv thaiv uas tau tsim los rov qab los ntawm NHR preinvasive thaum txoj hauv kev koom nrog cov txheej txheem ntau dua preinvasive tiv thaiv, suav nrog PEN2-txog txoj hauv kev, tsis muaj txiaj ntsig [ 48, 51] ib.
Qhov no yog raws li kev sib raug zoo ntawm A. thaliana thiab qis incompatible nonadapted pathogen C. nymphaea PL1-1-B, uas ib nrab overcomes lub high-txheej preinvasive tiv thaiv [51]; C. nymphaea PL1-1-B muaj zog ua rau ECR thiab tseem ua rau muaj kev tiv thaiv kab mob hauv cov nroj tsuag qus [48]. Ntau cov kab mob phytopathogenic fungi, xws li C. nymphaea PL1-1-B, uas ib nrab kov yeej cov txheej txheem kev tiv thaiv ntau dua ntawm A. thaliana yog suav tias muaj nyob hauv qhov xwm txheej. Yog li ntawd, cov nroj tsuag tau hloov kho ECR ua ib qho ntawm cov txheej txheem ntawm epidermal NHR tiv thaiv cov kab mob fungal no (Daim duab 4).
3. Epidermal Chloroplast-Localized Immune Cheebtsam Pab rau Preinvasive Antifungal Nonhost Resistance ntawm Arabidopsis
3.1. Qhov Kev Tshaj Lij Tshaj Lij Tshaj Lij Tshaj Lij Tshaj Lij Tshaj Lij Tshaj Tawm rau Motile Epidermal Chloroplasts hauv A. thaliana
Yuav ua li cas ECR pab txhawb rau preinvasive NHR tiv thaiv cov kab mob uas tsis muaj kab mob fungal tseem yuav tsum tau ua kom tiav. Fluorescence nyob cell imaging qhia tias cov tshuaj tiv thaiv kab mob, xws li -glutamylcysteine synthetase GSH1, Ca2 ntxiv -sensing receptor CAS, MATE tsev neeg thauj EDS5, isochorismate synthase ICS1 / SID2, thiab aminotransferase ALD1, tau nyob hauv thaj tsam ntawm epidermal me me chlorureoplass. [15,48,83–{13}}]. Qhov zoo siab, cov teeb liab fluorescent ntawm cov khoom no hauv epidermal chloroplasts muaj zog ntau dua li cov hauv mesophyll chloroplasts loj [48,83,85,86]. Hauv ECR-activated pavement cells tom qab inoculation nrog nonadapted C. fioriniae CC1, GSH1, EDS5, thiab CAS cov proteins tau pom nyob rau thaj tsam ntawm thaj chaw ua ke nrog cov motile epidermal chloroplasts, qhia txog kev sib txuas ceev ntawm ECR thiab intracellular positioning ntawm cov no. lub cev tiv thaiv kab mob [48]. Cov kev soj ntsuam no tuaj yeem ntxiv dag zog rau qhov tseem ceeb ntawm epidermal chloroplasts hauv cov nroj tsuag tiv thaiv kab mob
3.2. Epidermal Chloroplast-Localized Immune Cheebtsam thiab Preinvasive Defense hauv A. thaliana
GSH1 pab txhawb rau cov nroj tsuag tiv thaiv ntawm glutathione biosynthesis nyob rau hauv ob qho tib si sib xws thiab incompatible A. thaliana-pathogen interactions [15,87]. CAS muaj kev koom tes hauv kev hloov pauv Ca2 ntxiv rau kev taw qhia hauv chloroplasts thaum cog kev tiv thaiv kab mob [88], thiab CAS-targeted fungal effector ntawm Sclerotinia sclerotiorum cuam tshuam nrog SA signaling hauv cov nroj tsuag [89]. ICS1/SID2 thiab EDS5 yuav tsum muaj rau biosynthesis thiab thauj ntawm SA precursors, feem, thaum lub sij hawm cog kev tiv thaiv kab mob [85,90,91]. Qhov sib txuas nruj ntawm cov khoom tiv thaiv kab mob no thiab cov kab mob epidermal chloroplasts cuam tshuam lawv txoj kev koom tes hauv epidermal NHR tiv thaiv cov kab mob fungal uas ua rau ECR. Kev koom tes ntawm GSH1, CAS, EDS5, thiab ICS1/SID2 rau preinvasive NHR tiv thaiv cov kab mob uas tsis tau hloov pauv; C. nymphaea PL1-1-B pom tau tias muaj MAE ntau dua nyob rau hauv lub epidermis ntawm gsh1, cas, thiab eds5 ib tug mutants piv rau cov tsiaj qus-hom nroj tsuag [51], whereas zoo xws li cov teebmeem ntawm cov mutations ntawm MAE ntawm C. fioriniae CC1, C. siamense MAF1, thiab P. oryzae tau pom nyob rau hauv txhua qhov kev hloov pauv nrog cwj mem2 hloov pauv (Daim duab 2) [48].
Qhov nce MAE tus nqi ntawm ntau incompatible C. siamense COC4 thiab C. orbiculare 104-T tau lees paub nyob rau hauv pen2 mutant nroj tsuag nrog ntau yam kev hloov pauv hauv cov tshuaj tiv thaiv kab mob (Daim duab 2) [48]. Yog li ntawd, tus cwj mem 2- nyob ntawm qhov cuam tshuam ntawm cov noob hloov pauv no, uas encode epidermal chloroplast-localized proteins, cuam tshuam nrog ECR tshwm sim los ntawm txhua tus kab mob fungal. Txoj kev tshawb no qhia tias Arabidopsis epidermal hlwb siv ECR-centered tiv thaiv kab mob, nyob rau hauv intracellular repositioning ntawm lub cev tiv thaiv kab mob cov proteins yuav txhim khu antifungal NHR (Daim duab 4) [48]. Cov kab mob epidermal chloroplast-specific protein, ALD1, yog qhov tseem ceeb rau kev tiv thaiv kab mob hauv zos thiab cov kab mob kis tau los tiv thaiv ob qho tib si kab mob thiab kab mob avirulent, thiab nws siv nws cov txiaj ntsig los ntawm kev tswj hwm SA sacumulation [86,92–94]. Nyob rau hauv lub neej yav tom ntej, nws yuav tsum tau piav qhia seb ALD1 tseem ua haujlwm hauv preinvasive NHR tiv thaiv cov kab mob fungal.
4. Epidermal Chloroplasts Accumulate ntawm qhov Interface nrog Oomycete Pathogen thiab CHUP1 Yuav tsum tau nkag mus rau hauv Nicotiana benthamiana
4.1. Focal Accumulation ntawm Epidermal Chloroplasts ntawm Interface nrog Oomycete Pathogen hauv N. benthamiana
Lub luag hauj lwm ntawm epidermal chloroplasts teb rau oomycete pathogen P. infestans kuj tau kawm nyob rau hauv cov qauv solanaceous cog N. benthamiana [95–97]. Hauv seem no, kuv hais txog cov kev tshawb fawb tsis ntev los no ntawm cov kev hloov pauv ntawm epidermal chloroplast txav thiab kev koom tes ntawm CHUP1 hauv actinomycete kev tiv thaiv kab mob, txawm hais tias muaj qhov tsis meej meej txog kev hloov pauv lossis tsis zoo ntawm P. infestans rau N. benthamiana, uas qhia tsis meej txog hnub nyoog muaj feem cuam tshuam. rau P. infestans; cov nroj tsuag paub tab yog resistant, whereas hluas cov nroj tsuag yog susceptible [98].
Thaum lub sij hawm teb ntawm N. benthamiana rau P. infestans, epidermal chloroplasts qhia intracellular txav thiab sib sau ua ke ntawm qhov sib cuam tshuam nrog cov kab mob haustorium, kev kis kab mob tshwj xeeb hyphae, hauv BAK1- ywj pheej (Daim duab 5) [96] . Yog li, focal immunity mediated los ntawm epidermal chloroplasts tau npaj rau hauv N. benthamiana-P. kab mob pathosystem. Ib qho tshwm sim zoo sib xws tau pom nyob rau hauv ECR-activated hlwb ntawm A. thaliana thaum lub sij hawm cov tshuaj tiv thaiv kab mob; Txawm li cas los xij, feem ntau ntawm cov chloroplasts saum npoo tau tawg nyob rau hauv rooj plaub ntawm ECR thiab, tsawg kawg nyob rau hauv lub tshuab ntsuas, tsis koom nrog cov kab mob sib txuas [48]. Muab hais tias epidermal chloroplast ntawm N. benthamiana feem ntau yog kuaj pom ntawm thaj tsam ntawm cov pavement hlwb, tsis hais cov kab mob inoculation, cov txheej txheem ua haujlwm ntawm epidermal chloroplasts teb rau cov kab mob tawm tsam tsis yog tib yam ntawm A. thaliana thiab N. benthamiana.
Tsis tas li ntawd, CHUP1- hais txog ECR thiab focal tsub zuj zuj ntawm epidermal chloroplasts mus rau qhov chaw nkag mus rau fungal tej zaum yuav yog cov xwm txheej ywj pheej hauv A. thaliana vim hais tias cov kab mob sib txuas tshwj xeeb ntawm cov kab mob epidermal chloroplasts hauv N. benthamiana yog CHUP{{2} }independent (Daim duab 5) [97]. Nyob rau hauv cov ntsiab lus no, nws tseem yuav tau piav qhia seb puas yog ib feem ntawm cov epidermal chloroplasts ntawm A. thaliana, uas focally accumulate nyob rau hauv cov fungal nkag mus, muaj ib tug txawv lub luag hauj lwm los ntawm ECR-associated epidermal chloroplasts, raws li zoo raws li nyob rau hauv N. . benthamiana.
4.2. CHUP1-Dependent Callose Deposition ntawm Oomycete Penetration Site thaum lub sij hawm Epidermal Resistance ntawm N. benthamiana
Savage et al. qhia tias qhov chaw ntawm epidermal chloroplasts ntawm qhov sib txuas ntawm N. benthamiana thiab haustoria ntawm P. infestans tshwm sim ywj pheej ntawm CHUP1 [97], uas txawv me ntsis ntawm Arabidopsis ECR [48]. Txawm li cas los xij, lawv tau pom tias CHUP1 yuav tsum tau nkag mus rau hauv cov hlwb epidermal ntawm N. benthamiana tiv thaiv P. infestans [97]. Silencing los yog knockout ntawm ob CHUP1 alleles nyob rau hauv N. benthamiana txhim khu susceptibility rau P. infestans, whereas lub tsub zuj zuj ntawm epidermal chloroplasts nyob ib ncig ntawm haustoria tsis impaired nyob rau hauv lub CHUP1 knockout mutant (chup1) [97]. Arabidopsis chup1 mutant kuj pom tias txo qis NHR tiv thaiv MAE ntawm nonadapted fungal pathogens [48], qhia txog qhov tseem ceeb ntawm chloroplast-associated protein CHUP1 hauv epidermal nroj tsuag tiv thaiv ob qho tib si oomycete thiab fungal kab mob.
Interestingly, focal callose deposition ntawm haustoria ntawm P. infestans, uas yog ib qho ntawm cov lus teb tiv thaiv ntawm cov nroj tsuag, yog CHUP1-nyob, qhia tias chloroplast-associated protein CHUP1 pab txhawb rau epidermal tiv thaiv kab mob oomycete kab mob los ntawm kev txhim kho. Kev nkag mus tsis kam ntawm callose deposition (Daim duab 5) [97]. Cov lus nug tseem ceeb yog yuav ua li cas qhov kev sib txuas ntawm cov callose deposition thiab focal accumululation ntawm epidermal chloroplasts ntawm lub pathogen interface tshwm sim vim hais tias CHUP1 yog ib tug chloroplast-kawg protein. Hauv qhov no, Savage et al. pom tsis muaj kev sib raug zoo ntawm callose deposition thiab muaj chloroplasts nyob ib ncig ntawm lub haustorium [97]. Raws li qhov kev tshawb pom no, hauv A. thaliana-C. orbicular incompatible kev sib cuam tshuam, callose deposition tsis cuam tshuam nrog ECR ua kom; cov callose tau txaus tso rau ntawm qhov chaw ntawm cov kab mob nkag mus rau hauv cov nroj tsuag tsiaj qus, qhov twg ECR tsis tau ua haujlwm vim muaj kev ua haujlwm ua ntej ntawm cov txheej txheem siab tiv thaiv preinvasive [48]. CHUP1 noob knockout hauv N. benthamiana tsis cuam tshuam rau lwm cov txheej txheem tiv thaiv kab mob, xws li PAMP-triggered MAPK phosphorylation, constitutively active MEK2DD-induced hypersensitive response (HR)-zoo li cell tuag, thiab effector-triggered HR cell death [97].
Yog li ntawd, Savage et al. speculated tias nyob rau hauv lub CHUP1-knockout N. benthamiana, txo nyob rau hauv zus tau tej cov epidermal chloroplast-derived molecules, xws li ROS thiab SA precursors, plausibly ua rau qhov tsis zoo ntawm callose tsub zuj zuj nyob ib ncig ntawm lub haustorium. Lawv kuj tsis suav nrog qhov muaj peev xwm uas cov duab thaij duab tsis raug cuam tshuam txog kev koom tes ntawm epidermal chloroplast mobilization mus rau haustoria hauv focal callose deposition, vim hais tias cov chloroplasts tau txav deb ntawm haustoria tom qab callose deposition [97]. Kuv cia siab tias yav tom ntej kev tshawb fawb los qhia meej txog cov txheej txheem tswj hwm ntawm callose deposition ntawm cov kab mob sib txuas ntawm chloroplast-sociated CHUP1 hauv N. benthamiana, thiab muab piv nrog cov tshuaj tiv thaiv kab mob epidermal chloroplast hauv A. thaliana.
5. Dynamic Morphology ntawm Epidermal Chloroplasts thiab Inter-Organelle Interactions hauv Cog Kev Tiv Thaiv
5.1. Stromule Formation, Enlargement, thiab Cavity Formation
Mesophyll thiab epidermal chloroplasts loj hlob dynamic tubular extensions hu ua stromules, uas muaj stroma [99]. Nyob rau hauv epidermal hlwb ntawm N. benthamiana, stromules nce thaum lub sij hawm antiviral thiab antibacterial immunities, qhov twg effector-triggered HR cell tuag tshwm sim [95]. N. benthamiana kuj induces stromule tsim thaum lub sij hawm PAMP-triggered tiv thaiv lub oomycete INF1 ntxiv rau cov kab mob flg22 thiab fungal chitin nyob rau hauv ib tug BAK1-dependent yam [95,96]. Interestingly, stromule induction yog suppressed los ntawm P. infestans effector AVR3a [96]. Stromule induction hauv epidermal chloroplasts kuj tau pom thaum lub sij hawm tiv thaiv kab mob hauv A. thaliana [95]. Ua ke, cov kev soj ntsuam no qhia txog kev koom tes ntawm stromules hauv cov lus teb tiv thaiv cov nroj tsuag. Qhov tseem ceeb, chloroplast txav thiab kev sib txuas nrog lwm cov organelles, xws li nuclei, yog kho los ntawm stromules thaum cog kev tiv thaiv kab mob [95,100]. Ntxiv mus, stromules tuaj yeem thauj cov kev tiv thaiv cuam tshuam txog kev taw qhia molecule ROS thiab chloroplast tiv thaiv protein NRIP1 mus rau lub nucleus hauv N. benthamiana [95,101]. NRIP1 yog yuav tsum tau rau cov tshuaj tiv thaiv kab mob hauv N. benthamiana [102]. Cov tshuaj tiv thaiv kab mob tiv thaiv kab mob GSH1, EDS5, thiab CAS kuj tseem nyob hauv thaj chaw rau stromules txuas ntxiv los ntawm cov kab mob epidermal chloroplasts hauv A. thaliana, txawm hais tias cov teeb liab hauv zos ntawm thylakoid protein CAS hauv stromules tsis muaj zog dua li ntawm ob cov proteins thiab tej zaum yuav yog ib qho khoom cuav. kev hais lus ntawm kev hais lus [48]
Yog li, hauv cov kab mob epidermal, cov tshuaj tiv thaiv kab mob tuaj yeem hloov kho lawv qhov chaw los ntawm chloroplast txav thiab nthuav lawv qhov chaw mus rau sab nraud ntawm chloroplasts ntawm stromules thaum lub sij hawm tiv thaiv kab mob. Txawm li cas los xij, nws tseem muaj kev tsis sib haum xeeb txog seb puas muaj cov stromules muaj txiaj ntsig zoo ntawm cov tshuaj tiv thaiv kab mob. Caplan et al. qhia tias CHUP1 gene-silenced N. benthamiana thiab Arabidopsis chup1 mutants pom muaj zog stromules, uas tau pom nrog NRIP1-GFP fusion protein, thaum tsis muaj cov kab mob thiab txhawb nqa effector-triggered HR cell tuag thaum lub sij hawm tiv thaiv kab mob thiab kab mob tiv thaiv kab mob. [95]. Hauv kev sib piv, Savage et al. pom tau hais tias chup1 mutants ntawm ob qho tib si N. benthamiana thiab A. thaliana nthuav tawm txo cov stromules, uas tau pom nrog GFP, thaum tsis muaj kab mob [97].
Ntxiv mus, chup1 mutant ntawm N. benthamiana tuaj yeem ua rau muaj kev tsim cov stromule txaus thaum sib tw nrog P. infestans, txawm tias nws muaj kev cuam tshuam ntau ntxiv rau cov kab mob no [97]. Txo qis NHR preinvasive hauv Arabidopsis nrog chup1 kev hloov pauv tawm tsam MAE-hom fungal pathogens kuj tau tshaj tawm [48]. Cov kev tsis sib xws no tuaj yeem piav qhia hauv cov kev tshawb fawb yav tom ntej. Nws yog noteworthy tias, nyob rau hauv cov nroj tsuag tiv thaiv, stromule induction thiab chloroplast txav, xws li ECR thiab focal tsub zuj zuj ntawm cov kab mob sib txuas, muaj ntau dua tab sis txawv cov txheej xwm, vim hais tias yav tas los nyob ntawm lub PAMP signaling kinase BAK1, thaum lub tom kawg tsis. Yog li ntawd, cov kab mob sib txawv ntawm cov kab mob sib txawv tuaj yeem ua ke los ntawm cov kab mob epidermal chloroplast-centered system kom sib haum xeeb cov lus teb tiv thaiv.
Lub epidermal chloroplasts yog qhov tsawg dua li mesophyll chloroplasts. Txawm li cas los xij, intriguingly, loj epidermal chloroplasts nyob rau hauv ECR-activated hlwb yuav raug soj ntsuam, thiab cov pej xeem ntawm cov chloroplasts loj hlob maj mam nce thaum incubation nrog cov nonadapted fungal pathogens (Daim duab 6). Enlarged chloroplasts tsis tau pom nyob rau theem pib ntawm ECR. Kev tsim cov kab noj hniav hauv qhov loj ntawm cov kab mob epidermal chloroplasts tau lees paub (Daim duab 6). Qhov tseem ceeb ntawm lub cev ntawm cov kev hloov pauv no hauv morphology ntawm epidermal chloroplasts thaum lub sij hawm ECR yog tam sim no tsis meej, tab sis epidermal chloroplasts yuav txhim kho cov preinvasive NHR los ntawm cov xwm txheej ntxiv nyob rau theem kawg ntawm ECR. Xwb, cog organellar morphology tej zaum yuav cuam tshuam los ntawm pathogen effectors, vim hais tias nws tau raug tshaj tawm hais tias ib ntus qhia ntawm Colletotrichum effector proteins ua rau kom loj ntawm nuclei nyob rau hauv epidermal hlwb ntawm N. benthamiana [103]. Yog hais tias xws li organellar expansions yog nyob rau hauv lub pathogen kab mob tswv yim los ntawm cov nyhuv, nws yog raws li qhov tseeb hais tias ntau pathogen effectors nkag mus rau chloroplasts thiab lub hom phiaj chloroplast-localized proteins [46,47].
5.2. Perinuclear Clustering ntawm Epidermal Chloroplasts thiab Nuclear Movements
Chloroplast paub tias yog pawg nyob ib ncig ntawm lub nucleus rau kev sib txuas lus ntawm lub cev, qhov chaw kis ntawm cov teeb liab retrograde tshwm sim hauv kev teb rau ntau yam kev ntxhov siab ib puag ncig (37,38,104-106). Hauv kev tiv thaiv cov lus teb ntawm N. benthamiana, retrogradeROS signaling los ntawm perinuclear pawg chloroplasts rau lub nucleus los ntawm stromules tau tshaj tawm (95); tib yam tau hais tseg hauv A. thaliana (88,107,108). Interestingly, qhov poob ntawm CHUP1 noob ua rau muaj zog perinuclear pawg ntawm epidermal chloroplasts nyob rau hauv ob qho tib si N. bnethamiana thiab A. thaliana (97. Perinuclear pawg ntawm epidermal chloroplasts kuj tau pom thaum lub sij hawm Arabidopsis ECR tom qab pathogen inoculation los yog hauv 10 CH. Yog li, ob hom nroj tsuag no muaj cov txheej txheem kev tswj hwm zoo sib xws los tswj cov kab mob perinuclear ntawm cov kab mob epidermal chloroplasts, uas CHUP1 qhia qhov tsis zoo. epidermal chloroplasts nyob rau hauv chup1 mutant ntawm A. thaliana [48], ntau dua perinuclear pawg ntawm epidermal chloroplasts kuj yuav ua hauj lwm deteriorate. Raws li lub tswv yim no, CHUP1-knockout N. benthamiana yog ntau raug rau P7 infestans [9] .
Lub teeb-induced nuclear zog nyob rau hauv Arabidopsis mesophyll thiab epidermal pavement hlwb yog CHUP1-dependent [74,78]. CHUP1 tshwj xeeb hauv zos rau lub hnab ntawv ntawm chloroplasts, tab sis tsis yog nuclei, thiab tswj cov chloroplast txav; yog li ntawd, CHUP1-kev sib haum xeeb nyob rau hauv lub cev txav ntawm epidermal chloroplasts yog lub zog rau lub zog ntawm lub zog hauv kev teb rau lub teeb muaj zog [74,78]. Ib yam li ntawd, nuclei, ua ke nrog perinuclear pawg chloroplasts, tsiv mus rau thaj chaw saum npoo thaum lub sij hawm ECR hauv A. thaliana, uas yog nruj nreem txuas rau tus nqi ntawm CHUP1 proteins [48]. Lub perinuclear endoplasmic reticulum kuj repositions rau epidermal nto thaum ECR [48]. Hauv N. benthamiana, lub nucleus txav mus rau qhov chaw nkag ntawm oomycete P. infestans [96,109]. Qhov no kuj tau pom nyob rau hauv lwm cov nroj tsuag pathosystems siv fungal thiab oomycete pathogens [110–112]. Lub focal accumululation ntawm lub nucleus tsis cuam tshuam los ntawm kev nce perinuclear pawg ntawm epidermal chloroplasts hauv CHUP1-knockout N. benthamiana [97]. Muab hais tias qhov overexpression thiab depletion ntawm CHUP1 proteins muaj qhov cuam tshuam ntawm epidermal chloroplast txav thaum lub sij hawm Arabidopsis ECR, nws yuav nthuav kom txiav txim seb puas perinuclear pawg ntawm epidermal chloroplasts yog suppressed los ntawm CHUP1 overexpression nyob rau hauv N. benthamiana, thiab li no seb lub focal accumulation. nucleus ntawm P. infestans interface yog perturbed.
6. Cov lus xaus
Qhov tseem ceeb ntawm chloroplast muaj nuj nqi hauv cov nroj tsuag tiv thaiv kab mob yog dav lees paub. Kev tshawb fawb tsom mus rau hom chloroplast tam sim no tau tshwm sim hauv thaj tsam ntawm cov tshuaj tiv thaiv kab mob. Muab lub luag haujlwm ntawm cov nroj tsuag epidermis ua lub hauv paus ruaj khov los tawm tsam cov yeeb ncuab, xws li cov kab mob fungal thiab oomycete nrog cov peev xwm nkag mus ncaj qha, epidermal atypical chloroplasts yog qhov kev tshawb fawb zoo tshaj plaws vim tias lawv cov ntaub ntawv me me piv rau cov neeg paub zoo mesophyll chloroplasts. .
Tshwj xeeb, cov kev tshawb fawb tsis ntev los no tau qhia txog kev sib txuas ntawm cov kab mob hauv nruab nrog cev ntawm cov kab mob epidermal chloroplasts thiab cov nroj tsuag epidermal tiv thaiv kab mob, nyob rau hauv ECR, focal tsub zuj zuj ntawm epidermal chloroplasts mus rau lub pathogen interface, stromule tsim, inter-organelle kev sib txuas lus, thiab lwm yam txheej txheem tej zaum yuav orchestrated rau. txhim kho cov tshuaj tiv thaiv kab mob [48,95–97,113]. Chloroplast-localized CHUP1, uas yog Ameslikas pom tias yog tus tswj hwm ntawm chloroplast daim duab hloov chaw txav hauv cov hlwb mesophyll [76], muaj lub luag haujlwm tseem ceeb hauv cov xwm txheej epidermal chloroplast-hais lus teb rau cov kab mob. JAC1, tus tswj hwm ntawm photoinduced tsub zuj zuj teb ntawm chloroplasts, kuj koom nrog hauv qhov chaw ntawm lub cell ntawm epidermal chloroplasts thaum ECR [48]. Txhawm rau nkag siab txog cov txheej txheem hauv qab ntawm kev tiv thaiv kab mob los ntawm cov lus teb chloroplast-centered, xav tau kev tshuaj xyuas ntxiv txog qhov twg thiab seb cov khoom siv hauv chloroplast daim duab hloov chaw ua haujlwm li cas pab txhawb kev tiv thaiv cov lus teb hauv cov nroj tsuag epidermis.
Nyiaj txiag:
Qhov kev tshawb fawb no tau txais nyiaj los ntawm JSPS KAKENHI Grant Numbers 15K18648, 18K05643, thiab 21H02194, thiab los ntawm Thawj Coj Kev Tshawb Fawb Txog Cov Hluas (LEADER) txoj haujlwm ntawm MEXT.
Kev lees paub:
Kuv ua tsaug Yoshitaka Takano rau nws txoj kev pab thiab kev sib tham muaj txiaj ntsig.
Kev tsis sib haum xeeb ntawm kev txaus siab:
Tus sau tshaj tawm tsis muaj teeb meem ntawm kev txaus siab.
Cov ntaub ntawv
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