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1 !"#$%&'%()*%+*,,%-&,./*%01%02%/*/3452*%3&.26%&4752*,,*1%%

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

3 NO SIGNAL SEQUENCE! Cytoplasmic!!! NUCLEAR LOCALIZATION SEQUENCE!! Nuclear!!! MITOCHONDRIAL SIGNAL SEQUENCE!! Mitochondrial!!! ER SIGNAL SEQUENCE! Plasma membrane! Secreted! Resident ER and Golgi! Endosomes, Lysosomes!

4 The endoplasmic reticulum (ER) -- Largest membrane-bound organelle. -- Supply proteins to other organelles in the endomembrane system. -- ~1/3 of total cellular proteins. The secretory pathway ribosome ER Golgi cis medial trans PM 8M%

5 1960 s George Palade Imaging of the endomembrane system Electron Microscopy (EM)

6

7

8 Trafficking of GFP-VSVG Temperature shift from 40 o C to 32 o C Temperature sensitive vesicular stomatitis virus G protein (VSVG) mutant begins to traffic at permissive temperature. Endomembrane system

9 The endoplasmic reticulum (ER) Figure 12-8 Molecular Biology of the Cell ( Garland Science 2008) -- Distinct functions -- One continuous organelle -- Ratio varies across cell types

10 The rough ER site for protein synthesis (Lumen) Pancreatic acinar cell (Similar in other professional secretory cells) The smooth ER Leydig cell from testis -- Lipid synthesis -- Adapted to specific biological functions -- Detoxification (e.g. cytochrome p450 in liver )

11 Biochemical assays in cell-free systems Dounce homogenizer

12 Figure 12-37b Molecular Biology of the Cell ( Garland Science 2008)

13 Rough ER Smooth ER

14 Cell-free transport assay Organelle proteins Soluble lumenal protein Cytosol Organelle Lumen Lipid-anchored proteins Integral Membrane protein -- These types of proteins are found in all organelles. -- Peripheral membrane proteins are not shown.

15 How are proteins targeted to organelles? Soluble lumenal protein Cytosol Organelle Lumen Integral membrane protein Ribosome Signals must exist to direct proteins:!! 1. Signal sequences for translocation! 2. Sorting or retention signals!

16 Translocation across the ER membrane! Soluble! protein! Membrane! protein! trans-membrane! (TM) segment! Signal! sequence! NO SIGNAL SEQUENCE! Cytoplasmic!!! NUCLEAR LOCALIZATION SEQUENCE!! Nuclear!!! MITOCHONDRIAL SIGNAL SEQUENCE!! Mitochondrial!!! ER SIGNAL SEQUENCE! Plasma membrane! Secreted! Resident ER and Golgi! Endosomes, Lysosomes!

17 Signal sequence directs protein targeting Each organelle has its own signal sequences for targeting Table 12-3 Molecular Biology of the Cell ( Garland Science 2008) Figure 12-35a Molecular Biology of the Cell ( Garland Science 2008)

18 Figure 12-35b Molecular Biology of the Cell ( Garland Science 2008) ER-targeting signal sequences 8./52%94&021.,02%?4&5,3./02% K*51(%02-*4(51*%L,&27%94&6.+(M% :;<=:><<?<<;<<;<=@?A?;;;BCDEFF%% % :G=DHCIJ<<<CJJ;KJB>@DFF% % :<<N;C<C<<;@C;;GIJ;BJ:HFF%% How do you confirm an ER signal sequence?

19 Translocation across the ER membrane! Soluble! protein! Membrane! protein! trans-membrane! (TM) segment! Signal! sequence! Recognition of signal sequence on the ER

20 Translocon from bacteria

21 Biogenesis of soluble lumenal proteins Example: BiP Figure Molecular Biology of the Cell ( Garland Science 2008) Biogenesis of type I membrane protein Figure Molecular Biology of the Cell ( Garland Science 2008)

22 Topologies of ER proteins Type I! membrane! protein! cytosol! C! N! C! N! C! Membrane! lumen! N! C! N! _! Other topologies! Soluble! lumenal! protein! Juxtamembrane residues may influence protein orientation

23 Membrane proteins have different topologies in ER membrane! cytosol! ER! lumen! Type I! N! cleaved! signal! sequence! C! -cleavable signal seq.! (7-15 apolar residues)! -transmembrane anchor! stops translocation! Type II! N! C! Type III! C! +! +! _! -signal anchor! seq. (18-25 apolar)! -not cleaved! _! N! N! +! -reverse signal! anchor! +! _! _!?! +! +! +! +! C! The topology of a protein is maintained:

24 ER chaperones and folding enzymes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isulfide bonds are common among ER proteins!"#$%&'(( )&*+,-)%((.*#/%"0*%(1!2.3% Disulfide bonds stabilize proteins and promote folding

25 N-glycosylation site: N-X-S/T Figure Molecular Biology of the Cell ( Garland Science 2008) N-glycosylation in the ER

26 N-glycosylation stabilizes proteins and promotes folding Properly folded ER proteins can be transported to other organelles in the endomembrane system

27 Incorrectly folded proteins are retained in the ER Incorrectly folded proteins are eventually degraded in the cytosol Figure Molecular Biology of the Cell ( Garland Science 2008)

28 ER stress response/unfolded protein response Ron and Walter Vesicular transport

29 Donor compartment Acceptor compartment Vesicles have a size of ~60-90nm

30 Genetic analysis of the endomembrane system (Randy Schekman, Nobel Prize, 2013) WT Sec12 mutant Sec17 mutant Budding yeasts Genetics: RNA interference

31 Three types of coat proteins for vesicle budding Figure 13-4 Molecular Biology of the Cell ( Garland Science 2008)

32 Proteins destined for export are captured by COPII coats Figure Molecular Biology of the Cell ( Garland Science 2008)

33 Assembly of COPII coat

34 Figure 13-13d Molecular Biology of the Cell ( Garland Science 2008) Dissociation

35 Transport vesicles Lipid transfer protein

36 The Golgi

37 BiP protein (human) M%.D+/+N"".+%+++/""'"**%*:DD*:N(7N%N(0:+(`9/%#N(Na)('N%*00"):P()'%% bm%07,/9n"c,%*(*'+0(:""%d)p+7/),*)%7n-:"d'+0(%'7b):,/npp%:0dd,ann%% MeM%*DD7D,90PN%:0(((P7Df%",**0/".N+%7D.D*7"*"9%+(DDN7$"NN%7N,"9-):"P%% MgM%'P"7D:"(3%"(+)N.'00)%*,7"""0"9(%+:D'*(*D)0%+N-:+(((f%:N/++3:)(%% eym%n-*nn"7)(:%7$+((*:-:p%'n.*$1d+9%ddd7(d:n'd%:)'"npd+''%*n*d"d"+//%% hvm%p$p"'0*0*/%-9*(*:-/*7%+7'"d-**+)%.:+-'/7.d,%npdn+*:/:+%dd/:0:*0n+%% hbm%n((/7'0,d0%pp+nd*i)(%d*,/'(0),:%*"n"9(""np%"(n+/(:p:7%(:+n++$n#,%% YeM%+7+(0*7N((%N.7D+0,/)7%NN,7D)/P0-%/7"/:)P,7N%3DN9*(*',%+7D:)$++(7%% YgM%-:+7(0,,",%'(N,P0*Nf%*0:N)(0+'N%7"*:D(7()D%)D037):P)%'+7,**0*'.%% \YM%N):"*D-"**%:DD+D*'0:7%')*+*/9"9/%+D)P0(:D*D%+((D+//*:D%*7.*D"N**D%% bvm%0*b+*/$p:"%:0*:a"ddd%*+**0np,00%/d+9(/"(,,%,7(**:7"*gav<%% ER retrieval signal

38 Retrieval of ER-resident proteins The KDEL receptor cycle KDEL receptor! KDEL! Retention/retrieval sequence! Golgi! low ph! Transport! vesicles! ER! neutral ph!

39 Three types of coat proteins for vesicle budding Figure 13-4 Molecular Biology of the Cell ( Garland Science 2008)

40 COPI coated vesicles mediate Golgi-to-ER and intra-golgi transport

41

42 W1/0./%(*(4&T06*%%%%%%%%%%%%%%%:522&10651*%II%%%%%%%%%%%%%%%%%%%E.+,*&106*%609)&19)5(51*%%% Modification of oligosaccharides in the Golgi

43 O-linked glycosylation occurs in the Golgi or serine Figure Molecular Biology of the Cell ( Garland Science 2008) Transport through the Golgi

44 Exocytic pathway Donor compartment Acceptor compartment Vesicles have a size of ~60-90nm

45 Figure Molecular Biology of the Cell ( Garland Science 2008) Vesicle docking and fusion

46 What drives vesicle fusion? Plasma membrane Vesicle (Sollner et. al. Nature 1993) Lab of Jim Rothman SNAREs (Soluble N-ethylmaleimide sensitive factor attachment receptors) The SNARE cycle trans-snare complex v-snare t-snares cis-snare complex NSF!SNAP ATP

47 Exocytosis

48 Three types of coat proteins for vesicle budding Figure 13-4 Molecular Biology of the Cell ( Garland Science 2008) COPI coated vesicles mediate Golgi-to-ER and intra-golgi transport

49 Two types of exocytosis Extracellular Space Golgi Constitutive exocytosis All cell types Stimuli Regulated exocytosis Neurotransmitter release in neurons Insulin secretion from pancreas Neurotransmitter release in synaptic transmission Presynaptic Terminal [Ca 2+ ] Postsynaptic Terminal Wormbook

50 H*(52.1%(&T02% The endocytic pathway - Endocytosis

51 S,5()402%(401X*,0&2%

52 ?I?Y%01%&2,Z%'&.26%&2%% ()*%9,51/5%/*/3452*

53

54 Figure Molecular Biology of the Cell ( Garland Science 2008)

55 Figure Molecular Biology of the Cell ( Garland Science 2008) Figure Molecular Biology of the Cell ( Garland Science 2008)

56 The endocytic pathway - Lysosome

57 Figure Molecular Biology of the Cell ( Garland Science 2008)

58 Figure Molecular Biology of the Cell ( Garland Science 2008)

59 Transport through the Golgi

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