Lub Innate Immune Teb Ntawm Eusocial Hymenopterans Rau Viral Pathogen Challenge
May 04, 2023
Abstract
Nyob rau hauv xyoo tas los no, kab tshuaj tiv thaiv kab mob tau nthuav dav sai hauv kev tshawb fawb txaus siab, thiab cov ntaub ntawv muaj tau nthuav dav hauv hom. Kab tiv thaiv kab mob los ntawm ntau yam kev coj cwj pwm thiab kev tiv thaiv lub cev tiv thaiv kab mob. Qhov kev xav tau ntawm kev tiv thaiv kab mob yog qhov tseem ceeb tshwj xeeb rau cov kab eusocial; Nestmate sib thooj ua rau muaj kev sib kis thiab kis kab mob. Tsis tas li ntawd, eusociality cuam tshuam nrog kev sib koom ua ke ntawm ntau txhiab tus tib neeg nrog cov yam ntxwv txo qis kev hloov pauv caj ces, ua rau muaj kev cuam tshuam rau cov kab mob sib kis.
Txhawm rau tiv thaiv qhov no, lawv tau tsim ntau cov lus teb rau cov kab mob, suav nrog kev tiv thaiv tib neeg lub cev tiv thaiv kab mob, kev tiv thaiv kev sib raug zoo, thiab kev tso tawm ntawm cov tshuaj muaj zog glandular. Kev tiv thaiv kev sib raug zoo uas ua haujlwm los ntawm Hymenoptera tau raug tshuaj xyuas thaum qhov kev tshuaj xyuas muaj n; viruseseseseseseses tsim los rau peb cov kev paub txog kev tiv thaiv innate ntawm eusocial Hymenopterans rau cov kab mob pathogenic invaders. Peb sib cav tias qhov kev tshuaj xyuas no yog qhov tseem ceeb rau kev nkag siab txog Hymenopteran biology thiab tseem ceeb heev rau kev nyiam siv. Peb sib cav ntxiv tias qhov cuam tshuam ntawm eusocial Hymenopteran innate tiv thaiv kab mob yog nyob deb; lawv txoj kev vam meej yog ib qho ntawm ob qho kev poob nyiaj txiag ntau heev nyob rau hauv cov ntaub ntawv ntawm cov ntsaum ntxeem tau thiab nyiaj txiag tseem ceeb hauv cov ntaub ntawv ntawm zib mu Apis mellifera.
Ob qho tshuaj tiv thaiv kab mob thiab tib neeg kev tiv thaiv kab mob muaj nyob rau hauv immunology ceg ntawm biology, tab sis lawv tsuas yog kawm cov kab mob sib txawv. Kab immunology kawm txog kev tiv thaiv kab mob ntawm cov kab thiab yuav ua li cas nws teb rau cov kab mob sab nraud, thaum tib neeg kev tiv thaiv kab mob kawm txog kev tiv thaiv kab mob ntawm tib neeg thiab seb nws ua haujlwm li cas los tiv thaiv cov kab mob sab nraud. Txawm hais tias cov khoom tshawb fawb ntawm ob qhov sib txawv, muaj qee qhov kev sib txuas ntawm ob. Piv txwv li, qee qhov kev tshawb fawb tau tshwm sim hauv kab tshuaj tiv thaiv kab mob tuaj yeem muab kev tshoov siab thiab siv rau tib neeg kev tiv thaiv kab mob, thiab qee qhov kev ua tau zoo ntawm cov tshuaj tiv thaiv kab mob peptide muaj nyob hauv kab tshuaj tiv thaiv kab mob kuj tuaj yeem siv los tsim cov tshuaj tiv thaiv kab mob rau tib neeg lub cev. Tsis tas li ntawd, kab tshuaj tiv thaiv kab mob kuj tseem muab qee qhov piv txwv ntawm kev tshawb fawb txog kev sib cuam tshuam ntawm cov kab mob thiab lub cev tiv thaiv kab mob rau tib neeg kev tiv thaiv kab mob thiab muab qee cov ntaub ntawv pov thawj rau kev tshawb pom thiab kev txhim kho cov tshuaj tiv thaiv tib neeg tshiab. Hauv peb lub neej niaj hnub, peb yuav tsum txhim kho peb txoj kev tiv thaiv. Peb pom tias Cistanche tuaj yeem txhim kho kev tiv thaiv. Cistanche yog nplua nuj nyob rau hauv ntau yam tshuaj antioxidant, xws li vitamin C, carotenoids, thiab lwm yam. Cov khoom xyaw no tuaj yeem tshem tawm cov dawb radicals, txo oxidative kev nyuaj siab, thiab txhim kho kev tiv thaiv ntawm lub cev.
Nyem cistanche tubulosa cov txiaj ntsig
Ntsiab lus:
tiv thaiv kab mob, eusocial Hymenoptera, kab mob.
Cov kab mob kis kab mob yog ib qho tseem ceeb tshaj plaws thiab tsis tshua nkag siab txog kev tiv thaiv kab mob tiv thaiv kab mob eusocial (Lester li al. 2019). Cov kab mob hauv tsev neeg Solinviviridae yog cov yeeb ncuab ntuj tsim ntawm cov kab mob loj eusocial cov kab liab tuaj tua hluav taws Solenopsis invicta (Valles li al. 2004, Valles thiab Rivers 2019) thiab tawny vwm ntsaum Nylandria fulva (Hymenoptera: Formicidaet) (Valles). 2012). Kev taw qhia ntawm cov kab mob no mus rau cov kab mob kis tau yog thaj tsam ntawm kev tswj hwm kev noj qab haus huv. Nyob rau tib lub sijhawm, tus zib mu Apis mellifera (Hymenoptera: Apidae) yog plagued los ntawm colony collapse disorder nrog tus kab mob kis (Chejanovsky li al. 2014), thiab colony kev vam meej yog nyob ntawm lub peev xwm los teb cov kev cov nyom los ntawm cov kab mob no. Invasive Ant hom yog lub luag haujlwm rau ntau lab nyiaj daus las hauv kev poob txhua xyoo rau ntau yam kev lag luam (Lard li al. 2006, Adams 2019).
Hloov pauv, zib ntab muv yog ib qho ntawm cov kab tseem ceeb tshaj plaws ntawm kev pab nyiaj txiag, thiab kev tswj hwm lawv txoj kev noj qab haus huv hauv kev teb rau cov kab mob uas ua rau mob plague lawv yog qhov tseem ceeb rau kev txaus siab ntau (Genersch 2010). Qhov kev tshuaj xyuas no ua rau lub cev tsis ntev los no thiab loj hlob ntawm kev ua haujlwm piav qhia txog kev tiv thaiv kab mob ntawm cov kab eusocial, nrog rau qhov kev txaus siab tseem ceeb ntawm kev tiv thaiv kab mob hauv lub cev. Peb pib los ntawm luv luv piav qhia txog kev tiv thaiv kev tiv thaiv kev sib raug zoo ntawm qib colony, them nyiaj tshwj xeeb rau kev tiv thaiv colony-scale. Peb mam li nqaim lub peev xwm mus rau tus kheej theem, tsom mus rau lub cev tiv thaiv kab mob. Peb nthuav tawm ib qho kev sib sau ua ke ntawm cov ntaub ntawv ntau ntxiv thiab kev tshawb fawb ntawm eusocial innate lub cev tiv thaiv kab mob rau cov kab mob kis kab mob thiab muab qee cov lus nug tshiab los mus ua haujlwm yav tom ntej hauv thaj chaw zoo siab no.
Eusociality nyob rau hauv kab tau hloov zuj zus ntawm nws tus kheej ntau zaus, tshwj xeeb tshaj yog nyob rau hauv termites thiab Hymenoptera (Nowak li al. 2010, Quiñones thiab Pen 2017). Txawm hais tias muaj txiaj ntsig ntawm eusociality, ib puag ncig tsim los ntawm kev sib koom ua ke kuj tsim cov xwm txheej zoo rau cov kab mob thiab cov kab mob; Kev muaj cov khoom noj tas li, muaj qhov tsis txaus ntseeg, thiab qhov chaw tos txais ambient kub muab qhov chaw tshwj xeeb rau cov kab mob siab thiab kab mob cab (Feldhaar thiab Gross 2008). Kev sib cuag ntawm cov neeg nyob ze tuaj yeem ua rau muaj kev sib kis ntau ntxiv rau cov kab mob sib kis, thiab nce kev sib raug zoo rau cov qauv eusocial ua rau muaj kev cuam tshuam zoo sib xws rau cov kab mob (Baracchi et al. 2012b). Qhov kev sib xyaw ua ke ntawm cov yam ntxwv no muaj txiaj ntsig zoo rau cov kab mob sib kis (Quevillon 2018, Pinilla-Gallego li al. 2020). Rau cov laj thawj no, qhov tshwm sim ntawm eusocial colony qauv txhawb nqa qhov xav tau rau cov lus teb muaj zog rau ntau yam invaders, ob qho tib si pathogenic thiab parasitic. Eusocial kab ua tau raws li qhov kev sib tw no los ntawm ntau txoj hauv kev, los ntawm kev tiv thaiv kab mob hauv zej zog rau tib neeg, tshuaj lom neeg hauv lub cev tiv thaiv kab mob.
Vim hais tias invertebrates, nrog rau eusocial kab, tsis muaj qhov tseeb antibody-mediated tiv thaiv kab mob, lawv txoj kev vam meej yog heev cuam tshuam los ntawm lawv lub peev xwm ua hauj lwm raws li ib tug unit nyob rau hauv colony-theem tiv thaiv kab mob (kev tiv thaiv kev sib raug zoo), thiab nyob rau hauv tus kheej theem ntawm innate tiv thaiv kab mob. cov lus teb (Erler et al. 2011, Myllymäki and Rämet 2014, Palmer and Jiggins 2015). Thoob plaws hauv keeb kwm, evolutionary "kev sib tw caj npab" tau nthuav tawm ntawm cov kab kab thiab cov kab mob kis rau lawv (Erler et al. 2011). Kab teb rau cov kab mob ntawm qib immunological yog thaj tsam ntawm kev txaus siab uas tau kawm zoo hauv yoov tshaj cum thiab Dipteran qauv kab mob Drosophila (Diptera: Drosophilidae) (Palmer and Jiggins 2015). Kev txaus siab tsis ntev los no tsom rau kev tiv thaiv kab mob hauv kev sib raug zoo Hymenoptera qhia txog kev paub txog lawv qhov tseem ceeb rau tib neeg nyiam.
Superorganism-Level Social Immunity
Lo lus "kev tiv thaiv kev sib raug zoo" yog hais txog kev coj tus cwj pwm hloov thiab kev hloov kho uas ua rau tib neeg thiab pawg neeg tiv thaiv kab mob. Kev tiv thaiv kev sib raug zoo yog nyob ntawm kev koom tes ntawm pawg neeg kom zam, txo qis, thiab tshem tawm cov kab mob parasitic (Cremer li al. 2007, Cremer and Sixt 2009, Erler et al. 2011, Cremer et al. 2018). Cov kab kev sib raug zoo qhia txog qhov zoo tshaj plaws ntawm cov kev coj cwj pwm no, kev tiv thaiv kab mob thoob ntiaj teb. Cov tswv yim sib tw yuav yog thawj qhov kev tiv thaiv tiv thaiv kab mob hem thawj; Kev sib raug zoo kab poj niam kev yug me nyuam feem ntau yog monogamous thiab khub niam txiv hauv ib qho kev tshwm sim, yog li txo qis kev kis tus kab mob thiab cov kab mob cab mus rau thawj cov menyuam yaus thiab ua rau muaj peev xwm kis tau tus kab mob ntsug (Cremer et al. 2007). A. mellifera lub cev tiv thaiv kab mob rau cov neeg tawm tsam pib sab nraud ntawm thaj chaw nrog tshwj xeeb gatekeeper zib mu muv nyob rau hauv lub colony nkag mus tsis kam cov neeg muaj kab mob los yog parasitized nkag mus rau lub zes (Waddington thiab Rothenbuhler 1976, Nruas thiab Rothenbuhler 1985, Cremer et al.
Thaum thawj kab ntawm kev tiv thaiv kev tiv thaiv tsis txaus los tiv thaiv tus yeeb ncuab ntxeem tau, ib lub tswv yim ntawm lwm yam kev tiv thaiv kab mob hauv zej zog suav nrog kev tshem tawm cov kab mob parasite, kev kub taub hau rau cov kab mob tua kab mob, thiab kev tiv thaiv kev tiv thaiv ntawm cov resins uas txhawb nqa thiab ua kom cov tshuaj tiv thaiv kab mob kis tau. (Cremer et al. 2007). Kev kub taub hau hauv cov ntsaum (Cremer 2019) thiab zib ntab muv (Goblirsch li al. 2020) tuaj yeem ua rau cov kab mob thiab cov kab mob parasitic invaders zoo, ua rau tsis zoo lossis txo cov kev hem thawj los ntawm cov neeg invaders (Sugahara thiab Sakamoto 2009, Goblirsch li 02).
Antimicrobial peptides kuj tseem txhawb nqa dav rau kev tiv thaiv kev sib raug zoo thiab kev tiv thaiv tus kheej; zib mu muv venom muaj antimicrobial zog thaum siv rau lub cuticle thiab tso rau hauv lub zuag (Baracchi li al. 2011, Baracchi li al. 2012a, Moreau 2013, Simone-Finstrom 2017). Ib qho tshwm sim zoo sib xws tau pom nyob rau hauv cov kab tsuag uas xaiv, nyob rau hauv uas daim ntawv thov cuticular ntawm venom zoo nkaus li muaj kev cuam tshuam kev huv (Tragust li al. 2013, Simone-Finstrom 2017). Kev coj tus cwj pwm tsis huv, suav nrog kev lees paub thiab tshem tawm cov neeg laus muaj kab mob (Baracchi li al. 2012b) thiab kev tshem tawm cov kab mob ectoparasites thiab cov kab mob ua kom huv si ua kev tiv thaiv nyob rau hauv ntau yam ntawm foraging eusocial Hymenoptera nrog rau sab hnub poob zib mu A. mellifera (Bozicic). 1995), Neeg Esxias zib mu Apis kerana (Hymenoptera: Apidae) (Rath 1999), thiab nplooj txiav ntoo Acromyrmex subterraneus (Hymenoptera: Formicidae) thiab Acromyrmex octospinosus (Hymenoptera: Formicidae) (Richard2et 9).
Thaum superorganism-theem kev tiv thaiv kev sib raug zoo yog qhov muaj txiaj ntsig zoo thiab tseem ceeb hauv kev teb thawj zaug rau kev nkag mus rau cov kab mob, nws feem ntau tsis txaus los kov yeej cov kab mob pathogenicity thaum tsis muaj kev tiv thaiv kab mob hauv lub cev (Brutscher li al. 2015). Qhov tsis muaj qhov tseem ceeb ntawm cov histocompatibility complexes, T-receptors, los yog immunoglobulins yam ntxwv ntawm vertebrate lub cev tiv thaiv kab mob ua rau ib tug hnyav kev cia siab rau ib tug neeg-theem mechanical teeb meem rau cov kab mob thiab innate (tshuaj kho mob) tiv thaiv kab mob. Ntawm cov kev paub txog kev cuam tshuam hauv eusocial kab innate tiv thaiv kab mob rau cov kab mob yog lub cev tsis muaj zog (Imd), Tus Xov Tooj, RNAi, thiab Janus kinase (JAK / STAT) txoj kev taw qhia, nrog rau kev tso tawm ntawm cov tshuaj tiv thaiv kab mob peptides (AMPs) (Barribeau thiab. al. 2015, Brutscher et al. 2015, Brutscher and Flenniken 2015).
Immune Deficiency (Imd) Txoj Kev
Hauv eusocial Hymenoptera, kev koom tes ntawm kev tiv thaiv kab mob tsis muaj zog lossis Imd txoj hauv kev tiv thaiv kab mob yog qee yam cryptic, thiab feem ntau ntawm kev tshawb fawb txog kab Imd tsuas yog rau Drosophila. Txoj kev Imd plays lub luag haujlwm nyuaj hauv keeb kwm ntawm kab kab, ua raws li ob qho tib si kev tiv thaiv kab mob thiab kev tswj hwm tseem ceeb hauv kev txhim kho (Erler li al. 2011). Hauv kev tiv thaiv kab mob ntawm Drosophila, Imd yog lub luag haujlwm rau kev tswj hwm ntawm NF-kB protein Relish (Dushay li al. 1996, Myllymäki thiab Rämet 2014), thiab qhia txog feem ntau ntawm Drosophila antimicrobial peptides (AMPs) (Myllymäki et al. 2014).
Hauv Drosophila, kev ua kom txoj hauv kev Imd pib nrog kev lees paub ntawm cov kab mob microbial, ua kom yooj yim los ntawm kev tshawb nrhiav cov kab mob sib txuas nrog cov qauv (PAMPs) uas yog cov cim rau cov kab mob. Kev ua imd feem ntau cuam tshuam nrog cov kab mob kab mob, tshwj xeeb yog Gram-negative thiab qee cov kab mob Gram-zoo, raws li PGRP-LC (transmembrane receptor) nyiam khi rau mesodiaminopimelic-acid tshwj xeeb rau cov kab mob (Kaneko thiab Silverman 2005, Kleino thiab Silverman 2014, 2019). Nws ua raws li tus thawj tswj hwm hauv kev pib ntawm Imd txoj hauv kev kis kab mob, lossis hauv thaj chaw nruab nrab ntawm cov lus teb nyob rau hauv ntu anterior ntawm midgut hauv zos kab mob (Choe li al. 2002, Gottar li al. 2002, ZaidmanRémy li al. 2006 , Myllymäki et al. 2014). Ntawm cov kev lees paub ntau yam yog peptidoglycan, ib qho qauv ntawm ntau cov kab mob kab mob.
Curiously, ectopic qhia txog PGRP hauv Drosophila lub cev rog tuaj yeem qhib qhov kev qhia ntawm antimicrobial peptides thaum tsis muaj kab mob (Myllymäki et al. 2014). Kev koom tes ntawm Imd txoj kev sib txawv nyob ntawm tus kab mob tus kab mob; Imd plays lub luag haujlwm ntau dua hauv Drosophila tiv thaiv tus kab mob Sindbis thiab Cricket Paralysis Virus dua li Tus Xov Tooj, qhov uas Imd tsis tshwm sim los cuam tshuam hnyav rau kev tshem tawm ntawm Drosophila C Virus (Brutscher li al. 2015).
Kev ua haujlwm tsis ntev los no txhawb nqa qhov cuam tshuam ntawm Imd hauv kev teb rau cov kab mob. Kev ua haujlwm molecular tshawb xyuas cov kab mob tiv thaiv kab mob hauv Argentine Ant Linepithema humile (Hymenoptera: Formicidae) tau nthuav tawm cov lus teb rau ntau hom kab mob. L. humile transcriptomic tsom xam ntawm kev tiv thaiv kab mob rau cov kab mob Pseudomonas spp. (Pseudomonadales: Pseudomonadaceae) kab mob thiab L. humile virus 1, whereas tus zib muv kab mob Black Queen Cell Virus tsis elicit ib tug loj hloov nyob rau hauv lub cev txoj kev qhia (Lester li al. 2019). Relish, ib qho kev hloov pauv hauv txoj hauv kev Imd, tau raug cloned thiab ua los ntawm ob qho tib si Nothomyrmecia macrops (Hymenoptera: Formicidae) thiab ntau hom Myrmecia ant, thiab yuav muaj feem cuam tshuam rau kev tso tawm ntawm cov tshuaj tiv thaiv kab mob peptides (Schlüns thiab Crozier 2007, Schluns thiab Crozier 2007, Schluns thiab Crozier 2007. ). Meta-analysis ntawm transcription-theem teb nyob rau hauv zib mu A. mellifera nyob rau hauv Varroa mite vectored kab mob kis kab mob kev kho mob kuj qhia txawv kev qhia ntawm Imd noob iap2 thiab rel (Doublet et al. 2017).
Thaum eusocial kab kev koom tes ntawm Imd hauv kev tiv thaiv kab mob tiv thaiv kab mob tseem muaj kev cuam tshuam me ntsis, kev ua haujlwm tsis ntev los no hauv Drosophila tau coj mus rau kev txheeb xyuas ntawm dreidel, cov noob uas encodes cov protein ntau cuam tshuam hauv Imd suppression. Deidel tau raug hijacked los ntawm ntau kab DNA kab mob, suppressing Imd txoj kev ua kom nyob rau hauv tus tswv tsev tiv thaiv kab mob rau tus kab mob. Qhov no tig mus rau qhov kev tshawb pom tias kinase IKK thiab qhov kev hloov pauv ntawm NF-K yog qhov tseem ceeb rau kev tshem tawm ntawm Drosophila C tus kab mob thiab kab mob Cricket Paralysis (Imler 2019). Ob leeg ntawm cov kab mob no yog cov tswv cuab ntawm tsev neeg kis kab mob Dicistroviridae, uas muaj ntau tus kab mob kis tau los ntawm eusocial Hymenoptera koom nrog.
Tus Xov Tooj Pathway
Raws li yog cov ntaub ntawv nrog ntau txoj kev tiv thaiv kab mob, feem ntau ntawm kev ua haujlwm ntawm Kab Toll tau tsom mus rau Drosophila. Tus lej xov tooj yog ib txoj hauv kev uas tau txais kev cawmdim thoob plaws ntau yam thiab yog ib qho tseem ceeb rau ob qho tib si lub cev tiv thaiv kab mob thiab kev loj hlob ntawm cov kab mob vertebrates thiab kab (Evans li al. 2006). Tus xov tooj yog feem ntau cuam tshuam rau cov kab mob fungal thiab Gram-zoo tiv thaiv kab mob, tab sis cov pov thawj qhia tau hais tias cov kab mob tiv thaiv kab mob tuaj yeem tshwm sim los ntawm kev hu xov tooj thiab (Doublet li al. 2017, Rosales thiab Vonnie 2017, Lester et al. 2019, Baty et al. 2020). Hauv Drosophila, Kev Hu Xov Tooj yog pib los ntawm kev sib cais ntawm cytokine-zoo li transcription factor Spatzle thiab kev khi ntawm C-terminal fragment ntawm leucine-nplua nuj rov (LRR) ntawm Tus Xov Tooj (Weber et al. 2007, Lindsay and Wasserman 2014).
Thaum Spatzle khi rau Tus Xov Tooj LRR, Tus Xov Tooj Dimerizes thiab ua haujlwm. Tom qab ua kom tiav, Tus Xov Tooj / interleukin -1 receptor homology (TIR) domains dimerize, uas txhawb nqa kev sib txuas ntawm cov khoom siv protein MyD88 ntawm nws lub npe TIR. MyD88 tom qab ntawd khi lub adapter protein Tube thiab nrhiav cov protein kinase Pelle uas khi rau cov neeg tuag. Pelle recruitment ua rau autophosphorylation ntawm Pelle, uas induces degradation ntawm inhibitor Cactus. Degradation ntawm Cactus induces tso tawm ntawm transcription yam xws li Diff los yog Dorsal, uas yog ces translocated mus rau lub nucleus (Lindsay thiab Wasserman 2014).
Tus xov tooj tseem cuam tshuam rau hauv lub cev tiv thaiv kab mob ntawm eusocial Hymenoptera. Qhov tseem ceeb, muaj ob qhov plausible Spatzle orthologs tshwm sim nyob rau hauv zib mu genome (GB13503, GB15688) (Evans li al. 2006). Sequencing ntawm zib mu genome kuj tau nthuav tawm ob homologs ntawm Drosophila transcription factor Dorsal, thiab cov khoom siv intracellular Cactin, Pellino, TNF receptor-associated factor-2, thiab Tollip.
Tag nrho cov khoom no tau ntseeg tias ua lub luag haujlwm tseem ceeb hauv Txoj Kev Toll, thiab txhua yam tshwm sim muaj nyob rau hauv ob qho tib si Drosophila thiab zib muv hom (Evans li al. 2006, Doublet et al. 2017). Cov kev tshawb fawb kev hloov pauv tau cuam tshuam tus xov tooj hauv lub luag haujlwm nyuaj hauv cov lus teb kab mob hauv cov muv muv; Cov tub ntxhais hluas muv sim kis tus kab mob Israeli Acute Paralysis virus (IAPV) nthuav qhia ntau ntxiv ntawm cov noob caj noob ces thaum paub tab ntau dua, cov kab mob kis tau zoo tsis tau qhia txog kev cuam tshuam ntawm Tus Xov Tooj (Brutscher li al. 2015, Galbraith li al. 2015). Thaum cov tshuaj tiv thaiv kab mob tiv thaiv kab mob tseem tsis tau nkag siab, kev ua haujlwm tsis ntev los no hauv Argentine Ant L. humile kuj tau txhawb kev ntseeg tias cov txheej txheem tseem ceeb ntawm cov tshuaj tiv thaiv kab mob koom nrog hauv cov tshuaj tiv thaiv kab mob rau cov kab mob sib tw (Lester li al. 2019). Txoj kev hu xov tooj NF-kB homolog dorsal-1A tau transcriptionally induced nyob rau hauv cov neeg ua hauj lwm caste zib muv thaum parasitized los ntawm mite Varroa, xav hais tias tus xov tooj yog cuam tshuam nyob rau hauv teb rau Deformed Wing Virus (DWV) kab mob, raws li Varroa yog lub luag hauj lwm rau DWV kis mus rau zib muv (Doublet et al. 2017).
Lwm yam khoom ntawm Txoj Kev Hu Xov Tooj tau raug txheeb xyuas hauv eusocial Hymenoptera ib yam nkaus. Qhov kev hloov pauv ntawm Relish tau raug cloned thiab sequenced hauv kab ntsaum Nothomytmecia macros nrog rau ntau hom Myrmecia, (Schluns thiab Crozier 2009, Johansson et al. 2013, Lindsay and Wasserman 2014). Ib yam li ntawd, tus xov tooj hu xov tooj rau txoj kev Cheebtsam Tus Xov Tooj, Pelle, thiab Dorsal tau cloned thiab sequenced hauv Formica aquilegia (Hymenoptera: Formicidae) (Lindsay thiab Wasserman 2014).
Txoj kev hu xov tooj hu xov tooj kuj tau cuam tshuam rau hauv lub cev tiv thaiv kab mob ntawm cov ntsaum kab mob thoob ntiaj teb rau kev sib tw ntawm cov kab mob kis. Lester et al, 2019 siv RNA sequencing tsom xam los txheeb xyuas cov kab mob uas kis tus kab mob Hymenopterans. Cov kab mob tiv thaiv kab mob feem ntau cuam tshuam nrog kev paub zoo RNA tus kab mob kis kab mob (Deformed Wing Virus thiab Linepithema humile virus-1) yog peptide paub cov proteins muab rau tus xov tooj thiab Imd txoj hauv kev cuam tshuam Argentine ntsaum (Lester li al. 2019 ). Qhov no qhia tau hais tias muaj kev cog lus muaj zog rau kev koom tes zoo sib xws ntawm Txoj Kev Hu Xov Tooj hauv cov kab hluav taws xob liab tawm teb rau cov kab mob Solenopsis invicta, vim tias cov kab mob no kuj tseem muaj txiaj ntsig zoo, ib leeg-stranded RNA kab mob (Valles li al. 2004).
Antimicrobial Peptides
Antimicrobial peptides (AMPs) muaj ntau haiv neeg, kev txuag tau zoo, thiab cov txiaj ntsig tseem ceeb muaj nyob hauv ntau hom kab mob vertebrate thiab invertebrate tiv thaiv kab mob. AMPs muaj feem cuam tshuam rau hauv lub cev tiv thaiv kab mob tiv thaiv ntau pawg kab mob thiab ua rau cov kab mob microbial ntawm cov kab mob. Lawv feem ntau muaj 15-20 amino acids thiab raug cais raws li lawv cov amino acid qauv thiab muaj pes tsawg leeg (Wu li al. 2018). Thaum AMPs feem ntau cuam tshuam rau cov kab mob kab mob, qee qhov ua tau zoo hauv kev teb rau cov kab mob sib tw (Evans and Lopez 2004, Yi et al. 2014, Wu et al. 2018).
Ntawm cov no yog cecropins, uas yog thawj zaug txheeb xyuas thiab cais tawm ntawm hemolymph ntawm cecropia npauj Hyalophora cecropia (Lepidoptera: Saturniidae) los ntawm lub npe los ntawm (Hultmark li al. 1982). Thaum cecropins feem ntau cuam tshuam rau kev tiv thaiv kab mob Gram-positive thiab Gram-negative, cecropin P1 inhibits viral particle tso tawm thiab attenuates virally induced apoptosis (Schluns thiab Crozier 2009, Guo et al. 2014, Wu et al. 2018), kis kab mob. Cov txheej txheem ntawm kev txiav txim cuam tshuam nrog kev cuam tshuam ntawm lub hnab ntawv viral thiab qhia tias muaj kev cuam tshuam tseem ceeb tiv thaiv kab mob tawm (Schluns thiab Crozier 2009, Guo et al. 2014, Wu et al. 2018).
Ob peb AMPs tau raug txheeb xyuas hauv cov ntsaum; defensins tau lees paub los ntawm cloning thiab sequencing nyob rau hauv 25 formicine Ant hom thiab 2 Myrmica hom (Schluns thiab Crozier 2009). Ntawm cov noob caj noob ces uas tau txheeb xyuas AMP yog ib qho kev cai rau serine protease inhibitor uas tuaj yeem cuam tshuam rau kev tiv thaiv kab mob (Schluns thiab Crozier 2009). Ob lub AMPs uas zoo ib yam li hymenopteran thiab dua los ntawm zib mu A. mellifera tau raug txheeb xyuas nyob rau hauv cov hluav taws kub tuaj txawv teb chaws Solenopsis invicta (Tian li al. 2004). Ob lub AMPs no tau nthuav tawm ntau zog hauv S. invicta poj huab tais uas nyuam qhuav dhau los ntawm cov poj huab tais uas tsis muaj kev sib raug zoo, qhia txog kev tiv thaiv kab mob cuam tshuam nrog kev sib deev (Evans et al. 2006). AMPs kuj tau raug txheeb xyuas nyob rau hauv venom ntawm ib co ntsaum (Kuhn-Nentwig 2003). Ntawm cov no yog cov insulins, uas qhia qhov zoo sib xws rau melittin los ntawm A. mellifera. Melittin yog antimicrobial thiab hemolytic (Schluns thiab Crozier 2009).
Bombus hom thiab A. mellifera tau pom tias muaj cov ntawv luam ntawm AMP defensin noob (1 daim ntawv hauv Bombus thiab 2 daim ntawv luam hauv A. mellifera) (Barribeau et al. 2015). Thaum ib zaug ntxiv thiab hymenopteran cuam tshuam nrog cov kab mob tiv thaiv kab mob, cov neeg sau ntawv sau tseg tias qhov loj me ntawm cov ntaub so ntswg cuam tshuam nrog deflation tuaj yeem ua rau kev tso tawm ntawm cov kab mob hauv lub cev, ua rau kev tso tawm ntawm AMPs hauv cov lus teb (Tian li al. 2004). Ib qho tshwm sim zoo sib xws tuaj yeem koom nrog hauv cov kab mob tiv thaiv kab mob. Cov zib mu A. mellifera yog qhov tshwj xeeb hauv nws daim ntawv thov AMPs ua ib daim ntawv ntawm ob qho tib si kev sib raug zoo thiab molecular tiv thaiv kab mob; Royalizing yog kab defensin cais tawm los ntawm koob muaj koob muaj npe jelly ntawm zib muv thiab ua lub luag haujlwm hauv kev tiv thaiv thiab kev loj hlob (Bílikova li al. 2015). Nws yuav tsum raug sau tseg tias AMPs tau txheeb xyuas hauv kev sib raug zoo Hymenopterans tau cuam tshuam rau cov kab mob (Gram-positive thiab Gram-negative) thiab cov kab mob fungal (Wu li al. 2018). AMP kev koom tes hauv eusocial Hymenopteran kab mob tiv thaiv kab mob tsis tau sau tseg rau peb qhov kev paub, tab sis cov pov thawj qhia tias qhov no tuaj yeem yog thaj chaw zoo siab ntawm kev tshawb nrhiav (Lester li al. 2019).
RNAi
RNAi, los yog RNA cuam tshuam / ntsiag to, yog txoj kev tswj hwm kev tswj hwm thiab tiv thaiv txoj hauv kev uas Drosophila thiab lwm yam kab tiv thaiv kab mob. Txoj hauv kev RNAi ua tiav los ntawm ob kauj ruam tseem ceeb, pib, thiab ua tiav (Van Rij li al. 2006, Zambon et al. 2006). Hauv kev pib, ob-stranded RNAs (dsRNA) tau ua tiav los ntawm endoribonucleses Dicer1 lossis Dicer2 rau hauv me me dsRNA ntu nrog 3 'overhanging kawg. Cov txheej txheem, me me RNAs yog siv rau kev pib ntawm RNAi cov kauj ruam ua tiav (Zambon et al. 2006, Brutscher and Flenniken 2015, Brutscher et al. 2015). Tom qab pib, cov RNA me me tau muab tso rau hauv RISC, lossis RNA-induced Silenced complex. Qhov no suav nrog kev khi cov qauv strand rau Argonaut protein. Lub anti-sensitive strand ntawm cov me dsRNAs ces muab tso rau hauv RISC los ntawm protein r2d2 (Schwarz li al. 2004). Lub singular strand ntawm dsRNA koom ua ke rau hauv RISC los ntawm Argonaut protein binding tseem tshuav thiab ua raws li cov lus qhia los nrhiav cov mRNA ntxiv.
Qhov no yog hu ua tus taw qhia strand. Kev khi rau mRNAs hauv lub xov tooj ntawm tes yog qhov tseeb vim yog kev taw qhia los ntawm kev sib koom ua ke ntawm cov hlua khi rau RISC thiab lub hom phiaj. Tom qab khi, Argonaut pib thiab catalyzes lub cleavage ntawm lub hom phiaj mRNA, ua rau degradation. Cov txheej txheem zoo ib yam li thaum microRNAs siv los ua cov lus qhia hauv Argonaut, tab sis kev taw qhia miRNA tuaj yeem ua rau muaj kev cuam tshuam tsis zoo thiab kev puas tsuaj ntawm mRNAs loj dua (Schwarz li al. 2004). Thaum tseem ceeb heev rau kev tswj hwm ntawm ob peb endogenous mRNAs, RNAi khi thiab degradation kuj muaj feem xyuam rau kev tswj cov kab mob thiab tau txais txiaj ntsig zoo hauv ntau yam kab mob xws li cov nroj tsuag, invertebrates, thiab qee cov kab mob (Robbins li al. 2009).
Txoj kev siRNA tau cuam tshuam rau kev tiv thaiv kab mob kis kab mob hauv Drosophila thiab lwm yam kab. Tus kab mob Flockhouse tau pom tias muaj qhov tsis zoo tiv thaiv Drosophila hauv cov yoov qus (50 feem pua ciaj sia 15 hnub tom qab kis kab mob), qhov Dicer -2 cov yoov mutant pom 60 feem pua kev tuag 6 dpi thiab 95 feem pua ntawm 15 dpi (Wang et al. . 2006). Txoj kev siRNA yog kev txuag tau zoo thiab tseem cuam tshuam rau hauv kev sib raug zoo Hymenopteran kab mob tiv thaiv kab mob. Feem ntau ntawm cov kab mob kis mus rau kev sib raug zoo Hymenopteran zib mu A. mellifera yog cov kab mob RNA hauv tsev neeg Dicistroviridae thiab Iflaviridae (Niu et al. 2014). Kev sib sib zog nqus tau ua rau cov kab mob colony-collapse kev txom nyem cov cheeb tsam tau txheeb pom muaj ntau 21-22 nucleotide siRNAs nrog cov kab ke zoo sib xws nrog Israeli Acute Paralysis Virus (IAPV), Kashmir bee virus, Deformed Wing Virus (DWV) (Hunter et al. 2010), thiab A. mellifera rhabdoviruses-1 thiab -2 (McMenamin thiab Flenniken 2018). Kev txo qis ntawm tus kab mob viral titer ntawm IAPV thiab DWV tau tshwm sim, txhawb qhov kev lees paub tias txoj hauv kev siRNA cuam tshuam rau hauv cov kab mob tiv thaiv kab mob xws li Drosophila (Hunter et al. 2010). Cov txiaj ntsig zoo sib xws tau pom thaum Israeli thiab United States zib ntab muv thaj tsam tau sim sim nrog yim hom kab mob sib txawv (Chejanovsky li al. 2014).
Ib yam li ntawd, Florida thiab Pennsylvania honey bee colonies elicited tseem ceeb tiv thaiv los ntawm qhov tsis zoo ntawm IAPV los ntawm homologous dsRNA raug (Hunter et al. 2010). Kev nthuav qhia ntau ntxiv ntawm dicer thiab argonaute-2 tshwm sim nrog rau ob qho tib si kev sim tus kab mob (Brutscher li al. 2017) thiab rau cov kab mob Israeli Acute Paralysis virus (IAPV) hauv zib mu (Galbraith li al. 2015). A. cerana larvae kho nrog dsRNA tshwj xeeb rau Suav Sacbrood tus kab mob kuj ua rau txo qis ntawm tus kab mob titer ntawm tus kab mob tam sim no nyob rau hauv cov cheeb tsam kuaj (Brutscher li al. 2015, Brutscher thiab Flenniken 2015). Cov txiaj ntsig no muaj feem cuam tshuam rau kev txaus siab rau cov kab ntsaum kab mob teb rau cov kab mob kis kab mob, ntau yam kab mob kis kab mob kis kab mob, lossis yav dhau los, suav nrog cov tswv cuab ntawm tsev neeg kis kab mob Dicistroviridae (Lester li al. 2019, Baty li al. 2020).
JAK/STAT
Hauv Drosophila, JAK / STAT (Janus kinase / signal transducer thiab activator of transcription) txoj kev ligands suav nrog peb cytokine-zoo li proteins hu ua unpaired (nce), unpaired2 (upd2), thiab unpaired3 (upd3). Txhua yam yog induced los ntawm cov ntaub so ntswg qhov txhab. Upd yog induced los ntawm cov kab mob tiv thaiv kab mob, tab sis upd2 thiab upd3 yog induced raws li cov lus teb rau kab mob kab mob. Hauv kev teb rau cov kab mob ntxeem tau, txhua qhov ntawm cov molecules tshwj xeeb khi rau ib qho receptor tshwj xeeb hu ua Domeless. Hauv Drosophila, muaj ib qho JAK (Avadhanula li al. 2009) thiab ib qho STAT transcription factor, hopscotch, thiab Stat92E, feem.
Nyob rau hauv teb rau kev khi ntawm ib tug cytokine rau ib tug receptor, lub receptor dimerizes thiab activates JAK. Cov activated JAKs ces phosphorylate ib leeg nrog rau cov tyrosine residues ntawm tus receptor, uas tom qab ntawd ua raws li qhov chaw docking rau Src homology 2 domains ntawm STAT molecule. STATs kuj yog phosphorylated los ntawm JAKs, tso cai rau lawv kom dimerize thiab hloov mus rau lub nucleus los ua tus txhawb nqa ntawm lawv cov hom phiaj (Myllymäki thiab Rämet 2014).
JAK/STAT txoj hauv kev hauv kab tau raug kawm zoo tshaj plaws hauv Drosophila, uas yog plagued los ntawm ntau tus kab mob ntuj tsim. Raws li qhov xwm txheej ntawm kab teb rau lwm cov kab mob, Drosophila lub cev tiv thaiv kab mob rau cov kab mob kis tau cuam tshuam nrog kev sib tham ntawm ntau lub tshuab, suav nrog Tus Xov Tooj thiab JAK / STAT. Cov tshuaj tiv thaiv kab mob tiv thaiv kab mob tshwm sim los ntawm cov kab mob fungal thiab cov kab mob kab mob invaders yog txawv ntawm cov kab mob uas raug ntxias los ntawm cov kab mob, qhia txog cov lus teb tshwj xeeb rau cov neeg invaders lawv lub hom phiaj (Myllymäki thiab Rämet 2014). Nws kuj tau raug sau tseg tias ntau cov noob tau qhib los ntawm JAK / STAT txoj hauv kev teb rau tus kab mob Drosophila C, thiab cov kab mob kis tau thiab cov neeg tuag nce ntxiv nrog qhov tsis txaus ntawm txoj hauv kev no (Myllymäki thiab Rämet 2014).
Nws tau raug pom tias Drosophila qhia ntawm cytokines Upd2 thiab Upd3 tau mob siab rau tom qab kis kab mob Cricket Paralysis Virus (tsev neeg Dicistroviridae) (Lamiable thiab Imler 2014). Qhov no yog qhov cuam tshuam rau kev tiv thaiv kab mob hauv cov kab ntsaum kab mob thiab cov muv muv, vim tias ntau tus kab mob kis rau cov eusocial Hymenoptera yog cov tswv cuab ntawm Dicistroviridae. Cov kab hluav taws tua hluav taws Solenopsis invicta kab mob tau xub muab tso rau hauv tib tsev neeg kab mob (Valles li al. 2004) thiab tuaj yeem teb zoo ib yam li cov qauv kev kawm ntau dua Drosophila. Tsis tas li ntawd, txoj hauv kev siRNA thiab JAK / STAT tau koom nrog hauv kev sib tham nrog ib leeg hauv kev teb rau cov kab mob ntxeem tau (Niu 2015), thiab JAK / STAT tau cuam tshuam txog kev tiv thaiv kab mob hauv Hymenopteran Bombus terrestris, buff-tailed bumblebee.
Txhawm rau ntsuas qhov kev koom tes ntawm JAK / STAT hauv B. terrestris kab mob tiv thaiv kab mob, Niu, 2015 silenced JAK / STAT tseem ceeb tivthaiv Hop ua ntej inoculation nrog Israeli Acute Paralysis Virus (IAPV) thiab Slow Bee Paralysis Virus (SPBV). Txoj kev tshawb no tsis pom muaj kev sib raug zoo ntawm IAPV titer piv rau kev tswj hwm cov muv tsis muaj kab mob, tab sis muaj qhov nce ntxiv hauv SPBV tus kab mob titer ntawm ob hnub tom qab kis kab mob, qhia txog ib ntus rau JAK / STAT qhia txog kev tiv thaiv kab mob hauv B. terrestris (Niu 2015) (Supp Material [online nkaus xwb]).
Cov lus xaus
Lub cev tiv thaiv kab mob ntawm kev sib raug zoo Hymenoptera rau qhov kev sib tw ntawm cov kab mob kis kab mob uas kis rau lawv yog ib qho kev sib tw, ntau lub ntsiab lus uas nyuam qhuav pib tsis ntev los no. Raws li cov kev tshawb fawb zoo li no tau nce zuj zus, lub ntiajteb txawj nqus thiab qhov tseem ceeb ntawm kev tiv thaiv kab mob hauv lub cev rau kev noj qab haus huv biodiversity tau ua tiav. Eusocial Hymenoptera yog ecologically thiab economically tseem ceeb ob qho tib si nyob rau hauv detrimental thiab pab muaj peev xwm. Ntawm cov no, zib mu A. mellifera yog ib qho ntawm cov kab muaj txiaj ntsig zoo tshaj plaws hauv tib neeg kev ua liaj ua teb, pab nyiaj ntau lab nyiaj daus las txhua xyoo rau thoob ntiaj teb teev (Brutscher li al. 2015, Popovska Stojanov et al. 2021). Txawm hais tias lub luag haujlwm tseem ceeb ntawm cov zib ntab muv thiab kev txuag kev txuag lawv, nce mus txog 30 feem pua ntawm Tebchaws Meskas cov neeg nyob hauv lub tebchaws tau tuag txhua xyoo, nrog rau ntau qhov kev poob vim yog Colony Collapse disorder (CCD) (Brutscher li al. 2015, McMenamin thiab Genersch 2015, McMenamin thiab Flenniken 2018).
Raws li tus thawj coj pollinator no txuas ntxiv mus rau ravaged los ntawm tus kab mob viral, qhov tsim nyog rau kev tshawb fawb ntxiv ntawm lub cev tiv thaiv kab mob hauv lub cev ua kom pom tseeb. Nyob rau qhov kawg ntawm qhov kev txaus siab spectrum yog invasive ant hom. Invasive ants muaj kev vam meej tshwj xeeb, mus rau qhov ua rau muaj kev lag luam loj thiab biodiversity poob. Nrog rau qhov no hauv siab, kev tshawb nrhiav kom muaj kev nyab xeeb, muaj txiaj ntsig, tus tswv tsev tshwj xeeb, thiab tus kheej tshaj tawm cov tshuaj tiv thaiv kab mob lom neeg los tawm tsam cov ntsaum ntxeem tau coj mus rau kev txheeb xyuas ntawm ntau tus kab mob ntuj tsim yeeb ncuab uas suav tias yog cov tshuaj tiv thaiv kab mob lom. Txawm hais tias daim ntawv thov viral ntuj yeeb ncuab yog qhov chaw zoo siab ntawm kev tshawb fawb, kev nkag siab ntawm tus tswv tsev-tus neeg sawv cev kev sib cuam tshuam thiab kev tiv thaiv kab mob ntawm tus tswv tsev rau cov kab mob yog qhov tseem ceeb rau kev tswj hwm kev noj qab haus huv. Raws li qhov cuam tshuam ntawm cov kab mob kis kab mob kis rau cov kab eusocial tau tshwm sim, nthuav dav raws li kev nkag siab ntawm cov kab mob kis kab mob ua rau muaj feem cuam tshuam rau kev tshawb fawb thiab kev ua liaj ua teb.
Kev tiv thaiv kab mob cuam tshuam los ntawm cov kab mob sib ze muaj ntau yam sib txawv thiab qee qhov tsis sib xws thoob plaws taxa, thiab tag nrho qhov cuam tshuam ntawm cov lus teb no tsuas yog npaj rau tam sim no. Cov kab mob tuaj yeem cuam tshuam kev cuam tshuam ntawm qee txoj hauv kev, thiab lub cev tiv thaiv kab mob rau txawm tias cov yeeb ncuab ze ze ntawm tus kab mob tuaj yeem sib txawv heev nyob rau hauv taxa. Tsis tas li ntawd nthuav yog qhov kev soj ntsuam uas, tshwj xeeb tshaj yog nyob rau hauv cov ntsaum, qee txoj hauv kev zoo tiv thaiv qee yam kab mob tab sis tsis yog lwm tus; Qee cov noob tau muaj kev cuam tshuam zoo nrog kev sib tw kis kab mob hauv Argentine ntsaum, thaum lwm tus tau cuam tshuam tsis zoo. Cov kab mob tiv thaiv kab mob rau cov kab mob kis kab mob yog thaj chaw ntawm kev tshawb fawb uas tseem muab ntau txoj hauv kev rau kev tshawb fawb tshiab, tshwj xeeb tshaj yog hais txog kab mob raws li kev tswj hwm kab mob ntawm cov kab kab. Kev ua haujlwm tsis ntev los no ntxiv tau pom lub luag haujlwm ntawm kev kis tau RNAs (Maori li al. 2019), transgenerational immune priming (Amiri li al. 2020), thiab cua sov cov proteins (McMenamin li al. 2020) hauv kev sib raug zoo thiab innate tiv thaiv. Muaj undoubtedly ntau yuav tsum tau kawm txog lub cev tiv thaiv kab mob cov kab mob nyob rau hauv eusocial Hymenoptera; lub neej yav tom ntej yog ci ntsa iab rau thaj chaw zoo siab ntawm kev kawm no.
Kev lees paub
Qhov project no tau txais nyiaj los ntawm Texas A&M University Invasive Ant Research thiab Management Seed Grant program.
Cov ntaub ntawv Cited
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