Intertwined Thiab Finely Balanced: Endoplasmic Reticulum Morphology, Dynamics, Function, Thiab Kab Mob Part 6
Apr 10, 2024
Lub ER tuaj yeem txuas tau zoo rau microtubules, los ntawm cov lus qhia tubule, junctions, orlong qhov ntev ntawm tubule [187,232]. Kev sib txuas zoo li qub ntawm ER thiab microtubules tej zaum yuav tsim los ntawm cov proteins uas tau hais dhau los CLIMP-63 [15], p180 [16,194], REEP1 [17], thiab Sec61 [234].
Protein yog ib qho tseem ceeb ntawm cov khoom noj uas tsim nyog rau peb lub cev. Nws pab peb tsim thiab tswj lub cev cov ntaub so ntswg qauv, synthesize enzymes thiab cov tshuaj hormones, thiab lwm yam. Tab sis qhov koj yuav tsis paub yog tias nws tseem ceeb heev rau peb nco.
Proteins yog ib qho tseem ceeb units uas pleev xim neurons thiab neurotransmitters, uas yog cov qauv tseem ceeb hauv peb lub hlwb rau cov ntaub ntawv xa mus thiab nco cia. Yog li ntawd, cov proteins ua lub luag haujlwm tseem ceeb hauv lub hlwb.
Ntau cov kev tshawb fawb qhia tau hais tias noj cov protein kom txaus tuaj yeem txhim kho peb lub cim xeeb, ua kom peb lub hom phiaj thiab kev xav, thiab pab peb zoo dua siv thiab ua cov ntaub ntawv.
Protein tuaj yeem cuam tshuam rau peb lub siab thiab kev xav. Hauv peb lub hlwb, cov proteins tuaj yeem ua rau peb muaj kev txaus siab thiab kev xav sib npaug los ntawm kev tsim cov neurotransmitters xws li dopamine thiab norepinephrine.
Tsis tas li ntawd, cov protein tuaj yeem pab peb tswj lub cev noj qab haus huv, thiab txhim kho peb cov metabolism thiab lub cev tiv thaiv kab mob, yog li ua kom peb tuaj yeem ua kom pom tseeb thiab ua cov ntaub ntawv zoo dua.
Feem ntau, cov protein muaj feem cuam tshuam nrog kev nco, kev xav, thiab lwm yam. Yog li ntawd, peb yuav tsum tau xyuam xim rau kev noj cov protein ntau ntxiv kom ntseeg tau tias peb lub cev thiab lub hlwb tuaj yeem tau txais kev txhawb nqa thiab kev tiv thaiv zoo, kom siv tau peb cov kev txawj ntse thiab kev txawj ntse. Nws tuaj yeem pom tias peb yuav tsum txhim kho kev nco, thiab Cistanche deserticola tuaj yeem txhim kho kev nco, vim Cistanche deserticola tseem tuaj yeem tswj hwm qhov sib npaug ntawm cov neurotransmitters, xws li nce qib ntawm acetylcholine thiab kev loj hlob. Cov khoom no tseem ceeb heev rau kev nco thiab kev kawm. Tsis tas li ntawd, Cistanche deserticola kuj tseem tuaj yeem txhim kho cov ntshav khiav thiab txhawb nqa cov pa oxygen, uas tuaj yeem ua kom lub hlwb tau txais cov as-ham txaus thiab lub zog, yog li txhim kho lub hlwb tseem ceeb thiab kev ua siab ntev.

Nyem paub ntxiv los txhim kho kev nco
Cov kev sib txuas no tau pom zoo los tswj ER network nro thiab qhov chaw [235]. Nws kuj tseem muaj peev xwm hais tias ER txuas yuav ruaj khov microtubules, raws li tau pom rau p180 hauv axons [194].
Ib qho teeb meem nrog kev ntsuam xyuas qhov no yog qhov tshaj tawm ntawm cov kab mob microtubule-binding protein yuav ua rau muaj kev ruaj ntseg (xws li, [16,234]) uas tej zaum yuav tsis pom ntawm cov qib endogenous. Txawm li cas los xij, qhov muaj peev xwm sib cuam tshuam ntawm ER tubules thiab microtubules yuav yog qhov tseem ceeb rau kev txiav txim siab yav tom ntej, tsis yog tsawg kawg vim tias stabilizedmicrotubules yuav yog qhov zoo tshaj plaws rau kinesin-1 thauj lwm cov ER tubules.
3.1.4. ER Kev cuam tshuam nrog Actin Cytoskeleton
Hauv cov tsiaj hlwb, microtubules ua lub luag haujlwm tseem ceeb hauv kev hloov kho ER ntawm cov txheej txheem sib txawv uas tau piav qhia saum toj no, nyob rau hauv cov nroj tsuag, algae, thiab qee cov fungi, actin-based motility yog qhov tseem ceeb (saib hauv [236]).
Cytoplasmic streaming, myosin-drivendirectional ntws ntawm cytoplasm, kuj ua lub luag haujlwm hauv cov nroj tsuag cell ER dynamics [237,238] .Ntawm, ib qho tshiab ER thiab microtubule-associated compartment (EMAC) tau txheeb xyuas nyob rau hauv cov nroj tsuag uas tau muab tso rau ntawm microtubules, yog li ntawd. tiv thaiv qhov ntws [239]. Ib yam li ntawd, ruaj khov tubular ER junctions kuj tau pom los koom nrog microtubules [240].
Ib qho kev xav tsis thoob rau kev thauj mus los ntawm ER kuj tau pom hauv C. elegans nematodes, qhov uas poob ntawm ER thiab lub hnab ntawv nuclear-localized nesprin, ANC-1, ua rau tsis muaj kev tswj hwm ntawm ER, mitochondria, thiab lipid droplets vim yog quab yuam generated raws li tus wormcrawls [241]. Txawm li cas los xij, nws tseem tsis tau paub meej tias dab tsi cytoskeletal tivthaiv ANC-1 khi rau.
Hauv cov tsiaj hlwb, myosin-V-driven ER motility tuaj yeem ua rau muaj kev cuam tshuam ER luv luv (xws li, [211,242]). Tsis tas li ntawd, cov proteins filamin tuaj yeem khi ER rau actin hauv nasembryonic fibroblasts [243] thiab kev sib cuam tshuam ntawm ER thiab actin filaments ntawm myosin1c tuaj yeem ua kom cov ntawv ruaj khov [188].
Qib ntawm kev cia siab ntawm ER network ntawm actin cytoskeleton txawv ntawm hom cell, txij li depolymerizing actin filaments nrog cytochalasin tsis tshua muaj txiaj ntsig ntawm ER network faib hauv VERO hlwb [20], thaum nyob rau hauv epithelial-zoo li cellline Huh-7, actin depolymerization ua rau cov ntawv me me, tsawg dua ER ntawm cov cellperiphery thiab qhov tsis raug ER qhov chaw, nrog cov cheeb tsam loj ntawm cov hlwb tsis muaj organelle [188].
Tsis tas li ntawd, thaum microtubules yog depolymerized, qeeb, actin-dependentretraction ntawm ER ntawm lub cell center yog pom nyob rau hauv ib co [23,186,244] tab sis tsis yog tag nrho [20] cultured cell hom, qhia tau hais tias ER-actin kev cuam tshuam cuam tshuam ER morphology.Qhov tseem ceeb, super -kev daws teeb meem nyob-cell duab tau nthuav tawm microtubule-ywj siab ERtubule txuas ntxiv [38], uas tej zaum yuav zoo vim yog actin-based motility.
3.2. Network hloov pauv
Cov tubules, junctions, thiab cov nplooj ntawv uas ua rau endoplasmic reticulum hloov pauv lub sijhawm. Txawm tias cov ntsiab lus tsim los ntawm lub network uas tsis tau ua kom zoo dua qub tsis yog nyob ruaj khov tab sis theej oscillate nyob ib puag ncig txoj haujlwm. Muab cov complexmorphology thiab ceev ceev dynamics ntawm ER, quantifying ER motion yog ib tug tam sim no kev sib tw nyob rau hauv lub teb.Ib dav txais kev ntsuas ntawm dynamics yog qhov nruab nrab squared hloov chaw (MSD).
MSD yog ib qho kev ntsuas ntawm qhov deb npaum li cas ib qho khoom yuav tsum mus dhau lub sijhawm. Muab xam rau MSD, thawj qhov sib txawv ntawm txoj hauj lwm ntawm qhov khoom ntawm lub sij hawm pib thiab cov khoom tom qab tau pom.

Txhua khub ntawm cov ntsiab lus sib cais los ntawm tib lub sijhawm sib txawv (lossis lub sijhawm lag luam) raug txiav txim siab, thiab qhov sib txawv nruab nrab ntawm txoj haujlwm rau tag nrho cov khub no muab MSD ntawm lub sijhawm lag luam tshwj xeeb. Ua lej, MSD, h∆r2(τ)i, txhais tau tias:
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qhov twg x(t) thiab y(t) muab qhov chaw ntawm qhov khoom ntawm lub sijhawm t thiab τ yog lub sijhawm lag luam. Fitting apower txoj cai rau MSD tso cai rau cov lus tsa suab piav qhia tias yog diffusive, sub-diffusive, orsuper-diffusive. Diffusion, los yog Brownian tsab ntawv tsa suab, piav qhia txog cov txheej txheem xws li gradualmixing ntawm cheeb tsam ntawm siab thiab qis concentrations los tsim ib qho kev faib tawm. MSDs fordiffusive motion ua raws h∆r2(τ)i ∝ τ. Anomalous diffusion yog lo lus siv los piav qhia txog qhov uas MSD tsis yog diffusive. Super-diffusion piav qhia txog kev thauj mus los, xws li lub zog ntawm cov protein-tsav, thiab MSDs ua raws h∆r2(τ)i ∝ τ qhov twg 1 < < 2. Ntawm qhov tod tes, sub-diffusive tsab ntawv tsa suab yog txuam nrog cov lus tsa suab hauv ib puag ncig neeg coob andresults hauv MSDs nrog exponent <1.
Los ntawm kev taug qab cov ntsiab lus hauv ER network thiab suav cov MSDs ntawm cov kab no, cov ntaub ntawv hais txog kev hloov pauv ntawm tus kheej ntawm ER tau raug qhia meej. ER junctions tau pom tias txav mus rau qhov sib txawv, hauv microtubule-dependent yam [245]. Nocodazole kev kho mob txo tus naj npawb ntawm diffusive thiab super-diffusive junctions, qhia tau hais tias kev cuam tshuam nrog lub cev muaj zog proteinsmay yog qhov ua rau ntawm cov lus qhia junction. Tubules hauv ER network oscillatesub-diffusively feem ntau, nrog qhov nruab nrab MSD exponent ntawm 0.5 [25].
Cov txiaj ntsig no yog kev pom zoo nrog qhov kev kwv yees theoretical ntawm h∆r2(τ)i ∝ τ1/2 rau semi-flexible polymersoscillating thermally hauv nruab nrab viscous [246]. Cov txiaj ntsig no qhia tau hais tias qhov kev txav ntawm lub network raug txwv, uas zoo ib yam nrog kev txav mus los hauv ib puag ncig uas muaj neeg coob coob xws li cytoplasm. Tsis ntev los no, tag nrho cov contours ntawm ib leeg tubules kuj tau taug qab, uas ua rau muaj kev piav qhia ntxaws ntxiv ntawm ER tubule dynamics [27]. Qhov kev faib tawm ntawm qhov perpendiculardisplacements los ntawm qhov nruab nrab tubule qaum txoj hauj lwm tau pom rau cov ntsiab lus raws lub tubule contour.
Lub asymmetry, lossis skewness, ntawm cov kev faib tawm no tau suav. Qhov tseem ceeb skewness qhia active, tsav dynamics, whereas txoj hauj lwm nrog negligibleskewness piav cov ntsiab lus oscillating thermally, tsis muaj active influences. Feem ntau ntawm cov ntsiab lus oscillated thermally, txawm li cas los xij, ib feem peb ntawm cov ntsiab lus taug qab tau hloov pauv nquag. Dynamic constrictions thiab bulges nyob rau hauv txoj kab uas hla ntawm ER tubules kuj tau pom [38,181], txawm hais tias cov yam ntxwv ntawm cov oscillations tseem tsis tau soj ntsuam indepth. Raws li qhov tshwm sim ntawm ob qho tib si MSD thiab contour nrhiav kev tshawb fawb, ER tubules tau xav tias tooscillate sub-diffusively feem ntau, nrog rau kev ua kom nquag plias tshwm sim qee zaus raws lub tubule qaum. Lub sub-diffusive fluctuations yuav tshwm sim los ntawm cov lus tsa suab thiab cov neeg coob coob ntawm lub cytoplasm, whereas active dynamics tej zaum yuav tshwm sim los ntawm lub cev muaj zog proteins los yog membrane tiv tauj qhov chaw nrog trading organelles.
Sheet dynamics tsis tau kawm nyob rau hauv kom meej ntau npaum li tubular dynamics. Qhov no yog vim qhov nyuaj ntawm kev ntsuas qhov kev txav ntawm cov khoom lamellar, tshwj xeeb tshaj yog cov nyob hauv cov hlwb. Tshwj xeeb, cov cheeb tsam lamellar ntawm ER feem ntau nyob hauv qhov tuab ntawm lub xov tooj ntawm tes ze ntawm lub hauv paus, uas tawm ntawm lub teeb pom kev los ntawm lwm lub dav hlau ua rau kev ntes cov yeeb yaj kiab uas qhia tsuas yog ib txheej ntawm ER nplooj ntawv nyuaj. Txawm tias muaj cov yeeb yaj kiab pom tseeb uas qhia txog ER nplooj ntawv, qhov ntsuas qhov muaj peev xwm ntawm lamellar daim nyias nyias yog qhov nyuaj vim tsis muaj txoj hauv kev lees paub los piav qhia cov ntawv dynamics. Ob qhov kev ntsuas ntawm daim ntawv dynamics yog qhov nrawm ntawm daim ntawv npoo thiab qhov tawm ntawm lub dav hlau hloov pauv ntawm cov ntawv. Rau peb txoj kev paub, tsis muaj ob txoj hauv kev no tseem tsis tau siv rau cov ntawv ER perinuclear hauv vivo. Kev kwv yees theoretical tau ua rau kev tawm ntawm cov phiaj xwm ntawm cov nplooj ntawv membrane.
MSD ntawm cov undulations yog kwv yees kom nce asτ2/3 rau cov membranes dawb thiab qeeb qeeb, raws li ln(τ), rau cov membranes nyob rau hauv nro [246] .Kev sim, qee cov kev tshawb fawb tau txiav txim siab qhov dynamics ntawm lamellar ER ntawm lub cellperiphery. Joensu et al. [188] nrhiav pom tias qhov chaw thiab qhov muaj zog ntawm cov nplooj ntawv peripheralER yog nyob ntawm actin filament arrays thiab foci. ER nplooj ntawv tau pom los hloov pauv hloov pauv hauv cov lus teb rau kev txav ntawm cov qauv actin. Qhov ploj ntawm actin coj mus rau ER nplooj ntawv sau rau hauv qhov chaw seem tom qab thiab tsim cov qauv tshiab ua rau qhib lub fenestration hauv ib daim ntawv ER. Cov dynamics ntawm sheetedges kuj tau kawm.
Cov ntawv hloov pauv hauv thaj chaw me me thiab pom tias tsis muaj kev nyiam rau kev taw qhia hauv cov hlwb tsis kho. Kev kho mob nrog cov actin polymerization inhibitor, latrunculinA, nce qhov kev txav mus los ntawm cov nplooj ntawv raws li qhov kev faib ua feem ntawm cov nplooj ntawv undergoingfission, fusion, los yog transformations mus rau hauv tubules.Cov kev tshawb fawb ntxaws nyob rau hauv nqe lus no qhia tau hais tias qhov hloov pauv ntawm cov qauv tsim nyob rau hauv ER yog complex, varied. , thiab cuam tshuam los ntawm ntau subcellular organelles thiab txheej txheem. Kev sib cuam tshuam ntawm ER dynamics thiab dynamics ntawm lwm cov subcellularorganelles thiab cov qauv tsuas yog pib to taub thiab fruitful kev tshawb fawb nyob rau hauv cheeb tsam no yuav tsum tsis ntev.
3.3. Dynamics ntawm Membrane thiab Lumenal Cheebtsam
Cov txheej txheem ua los ntawm ER koom nrog ntau ntawm transmembrane thiab lumenal proteins, ntau yam txav mus nrhiav kev sib tham nrog cov neeg koom tes. Lub zog ntawm cov proteins no, nrog rau cov lipids tsim ER daim nyias nyias, tuaj yeem cuam tshuam rau tag nrho cov kev hloov pauv ntawm lub cev.
Cov txheej txheem fluorescence xws li kev taug qab ib leeg (SPT), fluorescence recoveryafter photobleaching (FRAP), thiab fluorescence correlation spectroscopy (FCS, reviewedin [247]) tau siv los ntsuas cov lipid thiab protein ntau. Khoos phis tawj simulationshave kuj tau siv los kawm txog qhov kev hloov pauv ntawm cov khoom siv kos rau hauv daim nyias nyias, raws li cov fluorescent probes siv los taug qab cov khoom subcellular tau xav tias yuav cuam tshuam cov dynamics. Highconcentrations ntawm fluorescent dye Rhodamine yog npaj los ua hydrodynamicdrag, txo qhov diffusion coefficient ntawm cov khoom txaus siab txog li 20% [248].

Ob peb daim nyias nyias tau paub los cuam tshuam cov kev hloov pauv ntawm cov khoom sib txuas thiab cov lumenal: lipid rafts, protein concentration, protein folding txheej xwm, cytoskeletal kev sib cuam tshuam, thiab daim nyias nyias. Lipid rafts yog cov thawj ntawm pawg lipids thiab cov proteins uas txav mus rau hauv lub bilayer [249]. Diffusion tau pom tias qeeb qeeb los ntawm ib qho ntawm ob qho hauv lipid rafts [250], thiab lipids thiab cov proteins tuaj yeem dhau los ua ntu zus rau cov rafts, uas tau pom tias muaj kev cuam tshuam, kev sib txawv ntawm kev sib txawv [251].
Qhov siab dua ntawm cov protein nyob rau hauv cov lipid bilayer kuj paub tias qeeb lateral diffusion [252], nrog rau kev simulations xaus lus tias lateral diffusion nyob rau hauv cov ntaub so ntswg heev yog afactor ntawm 5-10 qeeb dua nyob rau hauv dilute membranes [253]. FCS cov kev sim kuj pom tau tias qhov folding txheej txheem ntawm transmembrane proteins cuam tshuam rau lawv cov lus tsa suab nyob rau hauv ER membrane.Ob peb cov proteins tau soj ntsuam, tag nrho cov tau pom kom txav sub-diffusively [254].Qhov anomalous exponent ntawm unfolded VSVG tau pom tias qis dua qhov ntawd. ntawm nws foldedform.
Qhov no qhia txog qhov muaj kev cuam tshuam ntau dua ntawm cov proteins uas tau nthuav tawm. Kev khi ntawm calnexin, ib qho transmembrane chaperone protein, kom nthuav tawm VSVG ua rau muaj kev nce ntxiv ntawm cov anomalous exponent xws li cov lus tsa suab yog indistinguishable los ntawm foldedform. Cov txiaj ntsig no qhia tau hais tias calnexin tuaj yeem tiv thaiv kev tsim cov teeb meem tsis zoo ntawm cov proteins uas tsis tau nthuav tawm.Collisions ntawm cytoplasmic domains ntawm transmembrane proteins thiab cytoskeletal filaments kuj paub tias qeeb lateral txav nyob rau hauv lipid bilayers [255], raws li tau qhia rau transferrin receptor (TfR) ntawm plasma membrane. Nyob rau hauv ib txwm muaj, qeeb, kev txav ntawm TfR tau pom; Thaum actin tau depolymerized nroglatrunculin, dawb diffusion tau pom [256].
Photoactivation thwmsim nyob rau hauv luam yeeb leafepidermal hlwb tab sis pom tau hais tias transmembrane proteins nyob rau hauv lub ER nthuav tawm qeeb qeeb, diffusive dynamics thaum kho nrog latrunculin B nyob rau hauv kev sib piv rau lub active dynamicsobserved nyob rau hauv lub hlwb tsis kho [257]. Qhov no feem ntau yuav yog vim qhov kev hloov pauv ntawm myosin-tsav ntawm ER hauv cov hlwb cog (Tshooj 3.1.4). Lwm qhov piv txwv ntawm transmembrane proteindynamics tau hloov pauv los ntawm cytoskeletal kev sib cuam tshuam yog qhov kev txav ntawm ER tawm qhov chaw. ERESmove sub-diffusively raws ER tubules hauv microtubule-dependent yam [61,180].
Loweranomalous exponents thiab me me diffusion coefficients tau ntsuas thaum lub hlwb raug kho nrog nocodazole, qhia tias microtubular kev ua ub no txhawb ERES dynamics.In simulations, thov nro rau daim nyias nyias, raws li yuav tshwm sim nrog lub cev muaj zog kev ua si, nce lateral diffusion coefficients ntawm lipids nyob rau hauv lub bilayer, tsis muaj. Hloov pauv lawv cov exponents [258]. Cov anomalous exponents yog sub-diffusive, nrog tus nqi ntawm ~0.75 pom rau tag nrho cov membrane tensions. Cov dynamics kuj tau pom tias yog nyob ntawm cov kev taw qhia hauv computer simulations.
Deviations nyob rau hauv cov kev taw qhia perpendicular mus rau lub bilayer tau pom muaj constrained, whereas lateral motion nyob rau hauv lub dav hlau ntawm lub bilayer yog tsis [259].Tau ua ke, cov kev tshwm sim qhia tau hais tias lub zog ntawm daim nyias nyias lipids thiab transmembrane proteins yog complex thiab nyob ntawm seb muaj pes tsawg leeg thiab. Lub xeev ntawm lipidbilayer, thiab raws li kev sib cuam tshuam nrog cytoskeleton.Qhov kev hloov pauv ntawm cov substrates hauv lub lumen ntawm ER kuj tau ntsuas kev sim. Kev txhais lus ntawm cov proteins nyob rau hauv lub lumen ntawm ER yog thawj zaug tshawb nrhiav siv ntsuab fluorescent protein (GFP) hauv 1999 [260]. Qhov kev txav ntawm GFP hauv ER lumen tau pom tias qeeb qeeb dua hauv cytoplasm thiab hauv mitochondria.
Lub zog ntawm calreticulin, ib qho lumenal chaperone protein, tau pom nyob ntawm qhov chaw folding ntawm ER [261]. Hauv cov hlwb quiescent, calreticulin tau pom los ua piv txwv tag nrho ER, thaum qeeb qeeb diffusion coefficients tau pom nyob rau hauv cov hlwb metabolizing. Ib qho kev soj ntsuam ib leeg tau qhia tias ob qho tib si calreticulin thiab ER-targeted lumenal HaloTag proteins tsiv nrog qeeb qeeb ntawm ER junctions dua li hauv tubules [181]. Cov pejxeem ceev tau poob qis thaum ATP depletion, qhia tias ATP-nyob ntawm lub cev muaj zog protein-mediated dynamics ntawm ER tuaj yeem ua rau muaj kev cuam tshuam ntawm lumenal Cheebtsam. Qhov sib txawv ntawm qhov sib txawv ntawm cov tubules thiab junctions tsis tau pom zoo rau lub transmembrane chaperone calnexin.
Kev kho mob ntawm Cos-7 hlwb nrog latrunculin B ua rau kom nrawm dua lumenal protein dynamics, asdid tshem N-glycans los ntawm cov proteins txaus siab [262]. Txoj kev tshawb no, nrog rau cov kev sim siv TfR tau piav qhia saum toj no [256], qhia tias actin tuaj yeem ua lub luag haujlwm tseem ceeb ntawm kev txav ntawm cov proteins thiab lipids hauv lumen thiab daim nyias nyias ntawm ER.A kev sib txuas ntawm cov lus tsa suab ntawm ER thiab cov lus tsa suab. ntawm lumenal ormembrane-bound Cheebtsam tseem tsis tau ua.
Txawm li cas los xij, ntau qhov kev xav tau raug npaj. Georgiades thiab al. npaj siab tias cov oscillations ntawm ER tubules txhawb kev sib xyaw ntawm cov reactants hauv lub lumen, ua kom cov txheej txheem biosynthetic ua hauv ER [25]. Ib yam li ntawd, Stadler et al. qhia tias cov lus tsa suab ntawm ERES yog tshwm sim los ntawm ERnetwork hloov pauv, uas muaj zog microtubule-dependent [180].

Holcman et al. coj mus kom ze sib txawv thiab hais tias ATP-raws li peristalsis-zoo li contractions nyob rau hauv ERtubule diameters tsim active thauj ntawm lumenal Cheebtsam [181]. Nws yog qhov tseeb tias kev txav ntawm ER thiab cov kev hloov pauv ntawm ER-neeg nyob hauv cov khoom sib txuas, thiab nws xav tias qhov kev txav ntawm ER nce kev ua haujlwm ntawm cov txheej txheem cellular nqa tawm hauv lub cev, txawm tias qhov no tseem tsis tau lees paub.
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