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7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
1/20
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http://elife.elifesciences.org/http://dx.doi.org/10.7554/elife.00048http://dx.doi.org/10.7554/elife.00048.001http://creativecommons.org/licenses/by/3.0/http://creativecommons.org/licenses/by/3.0/http://creativecommons.org/licenses/by/3.0/http://creativecommons.org/licenses/by/3.0/http://dx.doi.org/10.7554/elife.00048.001http://dx.doi.org/10.7554/elife.00048http://creativecommons.org/licenses/by/3.0/http://www.elifesciences.org/the-journal/open-access/http://elife.elifesciences.org/7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
2/20
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http://dx.doi.org/10.7554/elife.00048http://dx.doi.org/10.7554/elife.00048.002http://dx.doi.org/10.7554/elife.00048.002http://dx.doi.org/10.7554/elife.000487/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
3/20
Djii flmimk|
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http://dx.doi.org/10.7554/elife.00048http://dx.doi.org/10.7554/elife.000487/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
4/20
Djii flmimk|
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http://dx.doi.org/10.7554/elife.00048http://dx.doi.org/10.7554/elife.00048.003http://dx.doi.org/10.7554/elife.00048.004http://dx.doi.org/10.7554/elife.00048.005http://dx.doi.org/10.7554/elife.00048.006http://dx.doi.org/10.7554/elife.00048.007http://dx.doi.org/10.7554/elife.00048.007http://dx.doi.org/10.7554/elife.00048.006http://dx.doi.org/10.7554/elife.00048.005http://dx.doi.org/10.7554/elife.00048.004http://dx.doi.org/10.7554/elife.00048.003http://dx.doi.org/10.7554/elife.000487/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
5/20
Djii flmimk|
Algglk jp oi% jIlcj 6>?61?8j>>>02% @ML8 ?>%;::0-jIlcj%>>>02 : mc 6>
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7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
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http://dx.doi.org/10.7554/elife.00048http://dx.doi.org/10.7554/elife.000487/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
7/20
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http://dx.doi.org/10.7554/elife.00048http://dx.doi.org/10.7554/elife.00048.012http://dx.doi.org/10.7554/elife.00048.013http://dx.doi.org/10.7554/elife.00048.014http://dx.doi.org/10.7554/elife.00048.014http://dx.doi.org/10.7554/elife.00048.013http://dx.doi.org/10.7554/elife.00048.012http://dx.doi.org/10.7554/elife.000487/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
10/20
Djii flmimk|
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7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
11/20
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tjetmv) pm dmepvmi pvoetdvl~plme% Vopbjv) T% ~mgfj jrdiztlsji| vjiljt me p{m gjoet m Lvj?'`j~je`je
~mtp'pvoetdvl~plmeoi vjkzioplme pm dm~j {lpb JV tpvjtt8
l% Vjkziopj` Lvj?'`j~je`jep gVEO `jdo| "VL@@, m o iovkj oe` blkbi| tjijdp tjp m gVEOt)
oii m {bldb ovj ~vj`ldpj` pm fj pvoetiopj` f| gjgfvoej'fmze` vlfmtmgjt op pbj JV% Pbj
gjdboeltpld jopzvjt m gVEO `jkvo`oplme le T% ~mgfj ovj blkbi| vjgleltdjep m VL@@
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ll% ^vmdjttlek mFl~? gVEO {lpble lpt 4 ZPV) ijo`lek pm imtt m lpt ~mi|O poli) {bldbdmzepjv'
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~vmdjttlek tpj~ vjtjgfij pbj lelploi Lvj?'`j~je`jep ezdijmi|pld tpj~ m VL@@ fzp Fl~? gVEO
pbje jtdo~jt zvpbjv `jdo|%
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lek `lsjvkjedj m tmizplmet pbop `ljvjep t~jdljt jsmisj` pm odbljsj pblt pota%
Pbj ~b|tlmimkldoi vmij m VL@@ le T% ~mgfjJV tpvjtt ovltjt vmg letzdljedljt le boe`ilek pbj ~vmpjle'mi`lek imo` le pbj JV izgje
Bmgjmtpotlt) pbjvjmvj) doe fj vjjtpofiltbj` le p{m ~vledl~oi {o|t8 ledvjotlek pbj do~odlp| m pbj
JV pm boe`ij pbj imo`mv `jdvjotlek pbj imo` pm gjjp pbj do~odlp|% Bjvj {j tbm{ pbop gVEO
`jdo| doe tjvsj ot pbj tmij gjoet m vjtmislek JV tpvjtt {lpbmzp pvoetdvl~plmeoi z~'vjkzioplme m
diottldoi Z^V povkjp kjejt% Pbj l`jeplj` pvoetdvl~pt povkjpj` mv VL@@ dmg~mtj o tzftjp m
gVEOt) oii jedm`lek ~vmpjlet pbop vjtl`j le mv pvosjvtj pbj tjdvjpmv| ~opb{o|% Fjlek pvoetiopj` f|
gjgfvoej'fmze` vlfmtmgjt) pbjtj gVEOt ovj pbjvjmvj le oe o~~vm~vlopj djiiziov imdoplme pm
gjjp odplsopj` Lvj?%Ot oii ~vmpjlet jedm`j` f| VL@@ povkjp gVEOt jepjv pbj JV izgje) pbjlv t|epbjtlt f| `jelplme
dmepvlfzpjt pm pbj fzv`je m pbj JV ~vmpjle mi`lek godblejv|% VL@@ pbjvjmvj bji~t vj`zdj pbj
~vmpjle'mi`lek imo`% Lp lt ijtt dijov bm{jsjv {bjpbjv tzdb vj`zdplme {mzi` bosj o gohmv lg~odp
Le`jj`) o foda'm'pbj'jesjim~j doidzioplme le`ldopjt pbop le T% ~mgfj VL@@ vj`zdjt pbj pmpoi ~vmpjle
leczr lepm pbj JV f| mei| ?:*) jsje ze`jv pbj tjsjvj JV tpvjtt dme`lplmet jr~imvj` bjvj jr~jvlgje
poii|% Pblt jtplgopj `jvlsjt vmg mzv vlfmtmgj mmp~vleplek `opo le emvgoi st JV'tpvjttj` djiit8 [j
tdmvj` pbj vjioplsj pvoetioplmeoi jekokjgjep m oii gVEOt jedm`lek ~vmpjlet `lt~io|lek tlkeo
tjqzjedjt mv pvoetgjgfvoej vjklmet pm jtplgopj czr m ej{i| t|epbjtlxj` ~mi|~j~pl`jt le pm pbj JV
oe` doidziopj` pbj lg~odp m VL@@ me pblt tjp "tjj Gopjvloit oe` gjpbm`t,% Lp lt `ldzip pm jesltlme
bm{ o gjvj ?:* vj`zdplme m fzia ~vmpjle czr lepm pbj JV {mzi` tzdj pm oiijslopj oe mpbjv{ltj
ijpboi JV tpvjtt%
Lp lt ~mttlfij pbop VL@@ ~vjjvjeploii| povkjpt ~vmpjlet pbop ovj ~ovpldziovi| `ldzip pm mi` oe`
bjedj glkbp bosj o `lt~vm~mvplmeoi lg~odp me pbj ~vmpjle mi`lek imo` le pbj JV izgje% Le`jj`)Lvj? go| fj imdoilxj` pm pbj sldlelp| m gVEOt jedm`lek tzdb ~vmpjlet f| lepjvodplmet m lpt JV
izgjeoi zemi`j` ~vmpjle tjetlek `mgole {lpb ~mvplmet m pbj eotdjep ~mi|~j~pl`j dbolet pbop
bosj jepjvj` pbj JV izgje) ot ~vjslmzti| ~vm~mtj` "Bmiilje oe` [jlttgoe) 6>>31 Vme) 6>>3,
Oe oipjveoplsj oe` emp gzpzoii| jrdiztlsj slj{ ~mtjt pbop VL@@ qzoilpoplsji| dboekjt pbj kjej jr~vjt
tlme ~vmij% Le tz~~mvp m pblt emplme) {j empldj pbop pbj ~m~zioplme m VL@@ povkjp gVEOt lt blkbi|
tjijdplsj% gVEOt jedm`lek ~vmpjlet lesmisj` le il~l` gjpofmiltg ovj blkbi| jevldbj` "4?* m VL@@
povkjp gVEO ot dmg~ovj` pm 3%;* le oii JV'povkjpj` gVEOt,% Gmvjmsjv) {j e` pbop VL@@ povkjpt
jedm`lek ~vmpjlet lesmisj` le tpjvmi gjpofmiltg ovj ~ovpldziovi| jevldbj` "?4* ot dmg~ovj` pm ?%4*,
Bm{ vj`zdj` tpjvmi t|epbjtlt {mzi` dmzepjvodp pbj pmrld jjdpt m JV tpvjtt vjgolet zedijov% Mej
http://dx.doi.org/10.7554/elife.00048http://dx.doi.org/10.7554/elife.000487/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
12/20
Djii flmimk|
Algglk jp oi% jIlcj 6>?61?8j>>>02% @ML8 ?>%;::0-jIlcj%>>>02 ?6 mc 6>
Vjtjovdb ovpldij
~mttlfij jr~ioeoplme {mzi` fj pbop JV tpvjtt ilglpt tpjvmi jrlp pbvmzkb sjtldziov pvoet~mvp oe` o dmg
~jetopmv| vj`zdplme le tpjvmi t|epbjtlt fjdmgjt lg~mvpoep pm tztpole fotld JV zedplmet) ~jvbo~t f|
golepolelek o~~vm~vlopj gjgfvoej czl`lp| "Elittme jp oi%) 6>>?1 Cjek jp oi%) 6>>4,% Le pblt {o|
VL@@ "oale pm mpbjv `jkvo`oplsj ~opb{o|t, dmzi` o`hztp fotld gjpofmild ~ovogjpjvt le pbj dji
"Fjveotdmel jp oi%) 6>?6,%
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jp oi%) 6>>>?6,% Mzv vjtzipt ~vmsl`j p{m zvpbjv ilejt mjsl`jedj le tz~~mvp m pblt slj{8 vtp) {j tbm{ pbop oe Lvj? VEotj odplsj tlpj gzpoep) Lvj?"B?>?2E,
lt zeofij pm tztpole djii kvm{pb me JV tpvjtt'le`zdlek gj`lo% Pblt gzpoplme {ot `jtlkej` pm fimda
dopoi|tlt {blij vjpolelek b|`vmkje'fme`lek lepjvodplmet m pbj oglem odl` tl`j dbole% Le`jj`
pbj jqzlsoijep tlekij oglem odl` tzftplpzplme le T% djvjsltloj Lvj?) Lvj?"B?>2>E,) vj`zdjt dopoi|pld
odplslp| f| 5?>:'mi`% Mpbjv{ltj) Lvj?"B?>2>E, lt le`ltplekzltbofij vmg {li` p|~j Lvj?) fmpb le lpt
milkmgjvlxoplme oe` tpvzdpzvoi ~vm~jvpljt ot `jpjvglej` f| dv|tpoiimkvo~b| "Amvjee|ab jp oi%
6>??,% Tjdme`) {j tbm{j` pbop dijosj` VL@@ povkjp gVEOt dovv| o 6 )4'd|dild ~bmt~bopj kvmz~
{bldb lt o ~vjvjqzltlpj mv pbj ilkoplme vjodplme "pVEO ilkotj, ztj` le pbj kjemgj'{l`j go~~lek
m gVEO je`t dvjopj` z~me JV tpvjtt% Lvj? oe` pVEO je`mezdijotj "oe` pbj Lvj?'ogli| gjgfj
VEotj I mze` le goggoiloe djiit, ovj pbj mei| d|pm~iotgld ezdijotjt aem{e pm ~vm`zdj tzdb
~vm`zdpt%
Le T% djvjsltloj) Lvj? bot o tlekij aem{e tzftpvopj) Bod? gVEO "El{o jp oi%) 6>>:,% Bod? oe`
RF^? gVEO bosj blkbi| dmetjvsj` oe` vjo`li| vjdmkelxofij tpjg imm~ tpvzdpzvjt pbop `jgovdopj
pbj t~ildj tlpjt% Dijosokj mddzvt op o zelsjvtoii| dmetjvsj` K oi{o|t mze` op ~mtlplme 4 le pbj
tjsje'fotj imm~ "Kmexoijx jp oi%) ??61 Mlao{o jp oi%) 6>?>,1 bm{jsjv) pbj
~mtlplme m pbj tdlttlij K le pbj imm~t lt emp dmetjvsj`% Pbjtj tpvzdpzvjt pbjvjmvj `m emp ~vmsl`j o
tpvzdpzvoii| ~ioztlfij jr~ioeoplme% Oipjveoplsji|) Lvj?t izgjeoi `mgole go| fjdmgj ~vjjvjeploii|
jekokj` {lpb eotdjep ~mi|~j~pl`j dbolet pbop `lt~io| blkbjv oelp| oe`-mv imekjv jr~mtj` ~j~pl`j
tjqzjedjt) pbjvjf| tjijdplek gVEOt dm'pvoetioplmeoii| f| vjdmkelxlek jopzvjt le pbj jedm`j` ~vmpjle
Pbj dmedj~p m Lvj? vjdvzlpgjep) {bjpbjv pbvmzkb lepjvodplmet slo pbj eotdjep dbole mv jijgjept le pbj
gVEO ~jv tj) lt tz~~mvpj` f| mzv e`lek pbop gzpoplme m mej dijosokj tlpj "ZKYD ZDYZ, le Kot6
gVEO klsjt vltj pm dijosokj op oipjveoplsj tlpjt% Pbjtj `opo le`ldopj pbop imdoi ~vmrlglp| vopbjv pboe pbj
VEO tjqzjedj tzvvmze`lek pbj lggj`lopj dijosokj tlpj go| kzl`j tzftpvopj tjijdplme%
Le mv`jv pm tpoflilxj pbj ~vlgov| Lvj? dijosokj ~vm`zdpt) {j ztj` gzpoep djiit lg~olvj` le jrmtmgj
zedplme dopoi|xlek 4 : VEO `jkvo`oplme "Oe`jvtme oe` ^ovajv) ?0,% Lepvlkzleki|) {j mftjvsj` pbop Kot6gVEO `jdo| le Tal6 gzpoep tpvolet {ot mftjvsj` mei|pvoetljepi|) ~joalek op 4> gle opjv JV tpvjtt le`zdplme% Pblt alejpld fjboslmv tzkkjtpt pbop dijovoedj
m Lvj?'dijosokj ~vm`zdpt f| Tal6 go| fj o vjqzlvjgjep mv dmeplezj` Lvj? odplslp|% Pbzt) lelploplme m
VL@@ f| Lvj? oe` zvpbjv `jdo| go| fj mfilkopmvli| dmz~ij`%
^vmdjttlek mFl~? gVEOPbj mftjvsoplme pbop T% ~mgfj Fl~? gVEO dboekjt tlxj z~me JV tpvjtt `opjt foda pm ?
7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
13/20
Djii flmimk|
Algglk jp oi% jIlcj 6>?61?8j>>>02% @ML8 ?>%;::0-jIlcj%>>>02 ?4 mc 6>
Vjtjovdb ovpldij
bjvj) Fl~? gVEO lt oitm dijosj` f| Lvj?) |jp jtdo~jt `jdo|% Pbj Lvj? dijosokj tlpj tbovjt pbj togj
jopzvjt `jtdvlfj` ofmsj mv VL@@ povkjp gVEOt% @zvlek JV tpvjtt'le`zdj` ~vmdjttlek) Fl~? gVEO
imtjt o ~mvplme m lpt 4 ZPV oe` ~mi|O poli% Pbj vjtziplek ~vmdjttj` pFl~? gVEO lt gmvj tpofij oe`
bjedj lt ~vjtjep op oe ledvjotj` tpjo`|'tpopj dmedjepvoplme% O ~ioztlfij jr~ioeoplme mv pbj ledvjotj`
tpoflilp| mpFl~? gVEO lt pbj imtt m oe VEO `jkvme imdopj` le pbj tjsjvj` ~mvplme m pbj 4 ZPV
pFl~? gVEO lt odplsji| pvoetiopj` {lpb lpt vlfmtmgj `jetlp| ~ovoiijilek lpt ledvjotj` ofze`oedj
Oipbmzkb ~mi|O polit ovj kjejvoii| ileaj` pm tpoflilp| oe` pvoetioplmeoi jdljed|) bltpmej gVEOt) {bldb
ilaj{ltj ioda ~mi|O polit) ~vmsl`j ~vjdj`jedj mv tzdb oe jrdj~plme pm pbj vzij "Govxiz jp oi%) 6>>2,
Bltpmej gVEOt pjvgleopj le o {jii'dmetjvsj` 4 tpjg'imm~ tpvzdpzvj) {bldb ~vmpjdpt vmg jrmezdijmi|pld `jkvo`oplme% ^vmpjlet fle`lek pbjvj zedplmet oale pm pbj ~mi|O fle`lek ~vmpjlet mze` me mpbj
gVEOt pm jeboedj bltpmej gVEO pvoetioplme f| imm~lek foda pm pbj : do~ tpvzdpzvj "Toedbjx oe`
Govxiz) 6>>6,%
[b| pFl~? gVEO jtdo~jt `jdo| vjgolet pm fj jr~imvj`% ^mttlfij jr~ioeoplmet lediz`j
pbj ~vjtjedj m tjdme`ov| tpvzdpzvj jijgjept% Le`jj`) {j e` o ~vj`ldpj` dmetjvsj` bolv~le
tpvzdpzvj op pbj 4 pjvglel m pFl~? VEOt le tmgj ttlme |jotpt "T% ~mgfj) T% mdpmt~mvzt oe`
T% dv|m~blizt,1 bm{jsjv ~vjilgleov| gzpoplmeoi oeoi|tlt olij` pm soil`opj lpt lg~mvpoedj mv gVEO
tpoflilp| le T% ~mgfj% Oe oipjveoplsj ~mttlflilp| lt pbop pbj 4 je` mpFl~? gVEO go| fj dmsoijepi|
gm`lj`% Tzdb gm`ldoplme {mzi` ejj` pm fj vjtpvldpj` pm pbj 6'MB) fjdoztj pbj 4'MB kvmz~ lt tpli
oddjttlfij mv gm`ldoplme f| VEO ilkotj "Clkzvj 4`,% Le pblt vjkov`) pFl~? gVEO {mzi` vjtjgfij
glVEOt) {bldb ovj 6'M'gjpb|iopj`) dmejvvlek vjtltpoedj pm jkvo`oplme "\z jp oi%) 6>>:,%
Mzv `opo tbm{ pbop Fl~? gVEO lt pbj mei| gVEO le T% ~mgfj le pbj jr~vjttj` kjemgj pbop lt
tzfhjdp pm pblt zelqzj vjkzioplme% [b| {mzi` tzdb o tlekziov gjdboeltg bosj jsmisj` jrdiztlsji| m
Fl~?= Cvmg {mva le mpbjv t~jdljt) Fl~? jgjvkjt ot pbj gmtp ~ijlmpvm~ldoii| lg~mvpoep oe` ~vjdltji|
dmepvmiij` JV dbo~jvmej "Kzim{ jp oi%) 6>>6,% Gmvjmsjv) Fl~? bmi`t o zelqzj ~mtlplme le T% ~mgfj
{bjvj lp lt ki|dmt|iopj` "^l`mzr oe` Ovgtpvmek) ?>:,% Mej go| t~jdziopj pbop JV tpvjtt
le T% ~mgfj) ot le mpbjv t~jdljt) jeboedjt pzvemsjv m ki|dmt|iopj` ~vmpjlet oe` pbop pbj vjkzioplme m
Fl~? gVEO lt fjejdloi pm lpt tpoflilp| f| dmg~jetoplek mv tzdb imtt "Fjveotdmel jp oi%) 6>?6,%
Dmediztlmet
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bosj fjje dmetjvsj`) fmpb `jpoli oe` kimfoi dmetjqzjedjt m `m{etpvjog ~vmdjttjt bosj fjje
o`o~pj` pm tjvsj `ljvjep ejj`t% Pbj Z^V le fmpb T% djvjsltoj oe` T% ~mgfj ziiit o d|pm~vmpjdplsj
vmij) |jp pbj gjdboeltgt m jrjdzplek pblt pota ovj m~~mtj`% Le T% djvjsltloj) mi`lek do~odlp| lt
ledvjotj` slo pvoetdvl~plmeoi z~'vjkzioplme1 le T% ~mgfj pbj mi`lek imo` lt `jdvjotj` oe` pbj JV lt
vjtpvzdpzvj`% Le gjpoxmoe djiit) fmpb gm`jt m Lvj? odplslp| ovj gjvkj`) oe` j~je`lek me dme`lplme oe`
djii p|~j doe tjvsj ljvjep ~zv~mtjt% VL@@ doe ~vmpjdp djiit f| vjgmslek gohmv tjdvjpmv| ~vmpjle imo`t
ot lp lt pbj dotj le letzile tjdvjplek djiit "Il~tme jp oi%) 6>>2,) mv lp doe tjvsj pm odplsopj o~m~mpmpld ~opb
{o|t) ot lp lt pbj dotj le djiit jr~jvljedlek ~vmimekj` oe` zeglplkopj` JV tpvjtt "Boe jp oi%) 6>>
7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
14/20
Djii flmimk|
Algglk jp oi% jIlcj 6>?61?8j>>>02% @ML8 ?>%;::0-jIlcj%>>>02 ?0 mc 6>
Vjtjovdb ovpldij
Gopjvloit oe` gjpbm`t
EmgjediopzvjPbj zelj` dmesjeplme ztj` le pblt goeztdvl~p mv kjejt oe` ~vmpjlet lt fotj` me pbj pvjopgjep f|
Oifjvpt) jp oi% le Gmijdziov Flmimk| m pbj Djii ":pb j`lplme) Kovioe` zfiltblek,) ~okj rrrll% Le fvlj) oi
kjejt ovj `jempj` le lpoildt {lpb o do~lpoilxj` vtp ijppjv% Gzpoep oiijijt ovj le`ldopj` f| o~~je`lek o
`jtdvl~pmv pm pbj kjej eogj% ^vmpjlet ovj le`ldopj` le Vmgoe ijppjvt%
Tpvolet) ~iotgl`t oe` kvm{pb dme`lplmetTpoe`ov` gj`lo oe` kjemgj jeklejjvlek gjpbm`t mv ttlme |jotp {jvj ztj` ot `jtdvlfj` ~vjslmzti|
"Gmvjem jp oi%) ?
7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
15/20
Djii flmimk|
Algglk jp oi% jIlcj 6>?61?8j>>>02% @ML8 ?>%;::0-jIlcj%>>>02 ?: mc 6>
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dzipzvjt {jvj `lizpj` oe` kvm{e zepli pbj| vjodbj` oe M@ m >%6:% JV'tpvjtt {ot le`zdj` {lpb pzel'
dog|dle ot le`ldopj`% Opjv 4 bv) tpvjttj` djiit {jvj {otbj` mzv plgjt {lpb ~vj'{ovgj` \JT: gj`lo
pm vjgmsj pbj `vzk) pbj dzipzvj {ot vjo`hztpj` pm oe M@ m >%6:) oe` ledzfopj` le o 60 {jii ~iopj
"? gi ~jv {jii, msjv 46 bv op 46D% Pbj M@ {ot gjotzvj` jsjv| ?> gle le o gldvm~iopj vjo jv "T|ejvk|
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me tmil` gj`lo "\J:T, oe` ledzfopj` mv 4 `o|t op 4>D% Dmimeljt {jvj dmzepj` vmg pbj `lizplmet
tjvljt% Zepvjopj` djiit tjvsj` ot o dmepvmi%
Fl~? gVEO boi'ilj gjotzvjgjepPm `jpjvglej pbj boi'ilj mFl~? gVEO) {j dmetpvzdpj` oe lepjkvoplsj ~iotgl` "~HA?02, dmepolelek
pbj m~je vjo`lek vogj oe` 4 ZPV mFl~? gVEO ze`jv pbj dmepvmi m o pbloglej'vjkziopj` Egp?
~vmgmpjv% Pbj dmetpvzdp lediz`j` pbj Egp? : ZPV% Djiit {jvj kvm{e le JGG dmg~ijpj gj`lo "{lpb
mzp oe| pbloglej, mv 60 bv% Djiit {jvj pbje `lizpj` lepm vjtb JGG dmg~ijpj gj`lo oe` vj'kvm{e pm
oe M@ m >%4% Opjv le`zdlek JV tpvjtt {lpb pzeldog|dle ">%6: k-gi, mv ? bv) pbloglej {ot o``j` pm
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http://dx.doi.org/10.7554/elife.00048http://dx.doi.org/10.7554/elife.00048.020http://dx.doi.org/10.7554/elife.00048.020http://dx.doi.org/10.7554/elife.000487/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
16/20
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7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
17/20
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7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
18/20
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7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
19/20
Djii flmimk|
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7/31/2019 The unfolded protein response in fission yeast modulates stability of select mRNAs to maintain protein homeostasis
20/20
Djii flmimk|Vjtjovdb ovpldij
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