The Signal Hypothesis and the Targe3ng of Nascent Polypep3des to the Secretory Pathway. Tuesday 8/31/2017 Mike Mueckler

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1 The Signal Hypothesis and the Targe3ng of Nascent Polypep3des to the Secretory Pathway Tuesday 8/31/2017 Mike Mueckler

2 Figure 6-63 Molecular Biology of the Cell ( Garland Science 2008) Ribosome Structure

3 Forma3on of Polyribosomes Figure 6-76 Molecular Biology of the Cell ( Garland Science 2008)

4 Intracellular Targe3ng of Nascent Polypep3des Default targe+ng occurs to the cytoplasm All other des+na+ons require a targe+ng sequence Major sor+ng step occurs at the level of free versus membrane-bound polysomes

5 Figure 12-36c Molecular Biology of the Cell ( Garland Science 2008)

6 Ribosomal Subunits are Shared Between Free and Membrane-Bound Polysomes Targe&ng informa&on resides in the Nascent polypep&de chain Figure 12-41a Molecular Biology of the Cell ( Garland Science 2008)

7 Signal-Mediated Targe3ng to the RER

8 Proper3es of Secretory Signal Sequences 8-12 Residues cleavage N ++ Hydrophobic Core Mature Protein Residues Located at N-terminus Residues in length Hydrophobic core of 8-12 residues OIen basic residues at N-terminus (Arg, Lys) No sequence similarity

9 In Vitro Transla3on/Transloca3on mrna Rough microsomes Ribosomes trnas System Soluble transla+on factors Low MW components Energy (ATP, crea+ne-p, crea+ne kinase) Re+culocyte or wheat germ lysate

10 Isola+on of Rough Microsomes by Density Gradient Centrifuga+on Figure 12-37b Molecular Biology of the Cell ( Garland Science 2008)

11 In Vitro Transla3on/Transloca3on System mrna + Transla+on Components + Amino acid* Protein* SDS PAGE

12 In Vitro Transla3on of Prolac3n mrna Prolac+n is a polypep+de hormone (MW ~ 22 kd) secreted by anterior pituitary MW (kd) SDS Gel Lanes: 1. Purified prolac+n 2. No RM 3. RM 4. No RM /digest with Protease 5. RM /digest with Protease 6. RM /detergent treat and add Protease 7. Prolac+n mrna minus SS + RM /digest with Protease 8. SS-globin mrna + RM / digest with Protease

13 Iden3fica3on of a Soluble RER Targe3ng Factor RM Centrifuge M KCl Supernate = KCl wash Pellet = KRM MW (kd) Lanes: 1. No addi+ons 2. KRM 3. KRM / digest with Protease 4. KRM + KCl wash 5. KRM + KCl wash / digest with Protease

14 Purifica3on of the Signal Recogni3on Par3cle (SRP) MW (kd) KCl Wash Hydrophobic Chromatography Lanes: SRP 1. No addi+ons 2. KRM 3. KRM /digest with Protease 4. KRM + KCl wash /digest with Protease 5. KRM + SRP /digest with Protease 8

15 Subcellular Distribu3on of the Signal Recogni3on Par3cle (SRP) Where is SRP located within the cell? 47% ribosomes + polyribosomes 15% cytoplasm 38% rough endoplasmic re+culum Conclusions: SRP likely moves between different subcellular compartments SRP is a soluble par+cle that can associate with membranes and is not a permanent membrane-bound RER receptor

16 Structure of the Signal Recogni3on Par3cle (7SL RNA) Figure 12-39a Molecular Biology of the Cell ( Garland Science 2008)

17 Interac3ons Between SRP and the Signal Sequence and Ribosome Figure 12-39b Molecular Biology of the Cell ( Garland Science 2008)

18 Iden3fica3on of an Integral Membrane Targe3ng Factor KRM Digest with Elastase Centrifuge E-supernate MW (kd) E-KRM pellet Lanes: 1. No addi+ons 2. SRP Only 3. SRP + KRM /digest with Protease 4. SRP + E-KRM 5. SRP + E-Supernate 6. SRP + E-KRM + E- Supernate

19 Iden3fica3on of SRP Receptor KRM Detergen t Solubilize SRP Affinity Column SRP Receptor MW (kd) Lanes: 1. No addi+ons 2. SRP 3. SRP + SRP Receptor 8

20 Structure of the RER Transloca3on Channel (Sec 61 Complex) Single-Pass 10 TMS Single-Pass Figure Molecular Biology of the Cell ( Garland Science 2008)

21 (Side-View) (From 2-D EM Images) (Lumenal View) Figure Molecular Biology of the Cell ( Garland Science 2008) A Single Ribosome Binds to a Sec61 Tetramer

22 Post-Transla3onal Transloca3on is Common in Yeast and Bacteria SecA ATPase func+ons like a piston pushing ~20 aa s into the channel per cycle Figure Molecular Biology of the Cell ( Garland Science 2008)

23 Classifica3on of Membrane Protein Topology Single-Pass, Bitopic Mul+pass, Polytopic

24 Genera3on of a Type I Single-Pass Topology Figure Molecular Biology of the Cell ( Garland Science 2008)

25 Type II Genera3on of Type II and Type III Single Pass Topologies Type III Post-transla+onal Transloca+on Figure Molecular Biology of the Cell ( Garland Science 2008)

26 Mul3pass Topologies are Generated by Mul3ple Internal Signal/Anchor Sequences Type IVa Figure Molecular Biology of the Cell ( Garland Science 2008)

27 Mul3pass Topologies are Generated by Mul3ple Internal Signal/Anchor Sequences + Type IVb Figure Molecular Biology of the Cell ( Garland Science 2008)

28 The Charge Difference Rule for Mul3spanning Membrane Proteins COOH NH COOH + NH 2 + cytoplasm NH COOH NH 2 + COOH + cytoplasm

29 NH 2 Transmembrane Charge Inversion Disrupts Local Membrane Topology in Mul3pass Proteins L1 L2 L3 COOH cytoplasm L NH 2 L2 L2 L3 COOH 2 3 NH L1 L2 L3 COOH 1 L1 L3 4 cytoplasm NH 2 COOH

30 N-Linked Oligosaccharides are Added to Nascent Polypep3des in the Lumen of the RER Figure Molecular Biology of the Cell ( Garland Science 2008)

31 Biosynthesis of the Dolichol-P Oligosaccharide Donor

32 Structure of the High-Mannose Core Oligosaccharide

33 Processing of the High-Mannose Core Oligosaccharide in the RER

34 Oligosaccharide Processing in the RER is Used for Quality Control Figure Molecular Biology of the Cell ( Garland Science 2008)

35 Disulfide Bridges are Formed in the RER by Protein Disulfide Isomerase (PDI)

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