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1 of 60 3 Gated trans Lectures 9-15 MBLG 2071 The n GATED TRANSPORT transport between cytoplasm and nucleus (bidirectional) controlled by the nuclear pore complex active transport for macro molecules e.g. nucleic acids & proteins active - costs the cell energy for transport nuclear envelope and nuclear pore complexes for them barrier and the gate nuclear import receptors are the adaptors between the gate and the cargo signalling sequences (nuclear localisation signals, NLS) identify the cargo Con The into Nu (NP env The Nuclear Envelope sport between cytosol nucleus Transport between cytosol and nucleus - continuous double and membrane (inner + outer) Gated transport Gated transport - the outer membrane extends into the The nuclear envelope endoplasmic reticulum Continuous double membrane Nuclear pore complex (NPC) cytoplasm Made up of approx. 30 proteins Nuclear Pore Complexes (NPCs) - perforate the nuclear envelope The outer membrane extends Aqueous pore that does not into the ER require protein unfolding (unlike - there is an average of 200 NPCs in mammalians transmembrane transport) - made up of approx. 30 proteins N u c l e a r p o r e c o m p l e x e s Gated transport - aqueous pore doesn'ttherequire (NPCs)that perforate nuclearprotein unfolding Tangles (like block the central envelope. opening for large molecules transmembrane transport does) Nuclear Localisation Signal (NLS) oplasm Bipartite signal patch - tangles block the central opening for large molecules (anything N above 40 kda can t get through in a significant number) Fig 12-8 Typically signal - free diffusion of smalla(<500 Da)patch molecules - limited diffusion (hence transport of folded proteins) of medium sized (<60 kda) molecules C eus Transport between cytosol and nucleus modified from: Nuclear Localisation Signal (NLS) NLS contain positively charged amino - typically a acids signal patch transport of folded proteins) lysine(hence and arginine - NLS contain positively charged amino acids, lysine and arginine SV40 Large T antigen Pro-Lys-Lys-Lys-Arg-Lys-Val Histone H2B Gly-Lys-Lys-Arg-Ser-Lys-Val Tumour suppressor p53 8 Lys-Arg-(aa)12-Lys-Lys-Lys ucleus eek-1.html nucleus nb: tumour suppressor p53 is often mutated in malignant tumour cases Nuclear Import Receptors - large family of proteins whose job it is to bind directly to a protein that needs to go into the nucleus, or via an adaptor protein which will then bind to transporting protein - i.e. several receptors that bind either NPC and NLS of cargo protein (A), or NPC and a nuclear import adaptor protein (B) [see image below]

2 6 of 60 Lectures 9-15 MBLG 2071 Co-translational translocation of proteins to the ER SRP = signal recognition particle Co-Translational Translocation Most proteins that are secretory, membrane-bound, or reside in the endoplasmic reticulum (ER), golgi or endosomes use the co-translational translocation pathway This process begins with the N-terminal signal peptide of the protein being recognized by a signal recognition particle (SRP) while the protein is still being synthesized on the ribosome The synthesis pauses while the ribosome-protein complex is transferred to an SRP receptor on the ER (in eukaryotes) There, the nascent protein is inserted into the translocon, a membrane-bound protein conducting channel (composed of the Sec61 translocation complex in eukaryotes) In secretory proteins and type I transmembrane proteins, the signal sequence is immediately cleaved from the nascent polypeptide once it has been translocated into the membrane of the ER (eukaryotes) by signal peptidase. The signal sequence of type II membrane proteins and some polytopic membrane proteins are not cleaved off and therefore are referred to as signal anchor sequences. Within the ER, the protein is first covered by a chaperone protein to protect it from the high concentration of other proteins in the ER, giving it time to fold correctly. Once folded, the protein is modified as needed (for example, by glycosylation), then transported to the Golgi for further processing and goes to its target organelles or is retained in the ER by various ER retention mechanisms. Post Translational Translocation Three distinct scenarios for PTT into the ER: a) soluble proteins b) proteins with a single pass transmembrane domain (i.e. can only cross membrane once) c) proteins with multi pass transmembrane domains (i.e. can cross several times)

3 16 of 60 Lectures 9-15 MBLG 2071 * highly processive (high ability to remain associated with template strand) * DNA pol ε synthesises the leading strand * DNA pol δ synthesises the lagging strand nb: there are many more DNA polymerases in eukaryotic replication however we will just focus on these DNA pol δ and ε Finishing Eukaryotic Replication eukaryotes face the problem of how to replicate the ends of linear chromosomes = END REPLICATION PROBLEM this is not a problem for prokaryotes as they have circular chromosomes see picture: incompletely replicated DNA - this is where the last RNA primer was and the last Okazaki fragment. There is no way you can fill in this gap with normal DNA pol machinery, but if this gap is left, and replication occurs again (in daughter chromosomes), then the chromosome would just get shorter, and shorter again for the next generation, etc Telomeres replication Telomeres solve end replication problem recall that the ends of eukaryotic chromosomes are composed of repeated structures called Telomeres in humans, the repeating sequence is 5 -TTAGGG-3 (no need to remember this) human telomeres can extend for as much as 10 kb a telomere is double stranded for most of it, but there is a short single stranded section where 3 end extends beyond 5 end end

4 22 of 60 Lectures 9-15 MBLG 2071 complex regulatory systems are in place to ensure orderly progression of the cell cycle A novel price in physiology of medicine was awarded in 2001 to the discoverers of these regulatory systems (Leland Hartwell, Paul Nurse, Tim Hunt) These prize winners used genetic and biochemical methods to identify the two classes of proteins that are key players in controlling the cell cycle Yeast as a Model Organism Hartwell worked on saccharomyces cerevisae daughter cell grows as bud from mother cell (i.e. the bud will be the daughter cell coming off the parent cell) followed progress of cell by observing bus growth Nurse worked on schizosaccharomyces bombe grows longitudinally before dividing followed progress of cell cycle by observing length of the cell Defining Mechanisms of Cell Cycle Control identify genes that regulate progression through cell cycle but if such a gene is mutated, the cell cycle will be blocked and this will most likely be lethal (and a dead yeast cell will not be helpful in the lab) circumvent this problem using conditional control conditional mutants display mutant phenotype under one growth condition but are normal when grown under another condition Look for yeast that show cell cycle arrest (i.e. stop copying DNA and dividing) at high temperature (36 o C) but are normal at 23 o C Temperature Sensitive Mutants Cell Cycle Mutants - CDCs; Cdks cell division cycle mutants = CDC mutants they are yeast cells that are stuck as particular stages of the cell cycle wee mutants on the other hand, speed up cell cycle cdc and wee genes later found to encode kinases and phosphatases that play critical roles in regulating specific stages of cell cycle 23 C RIP 36 C permissive temperature restrictive temperature

5 23 of 60 Lectures 9-15 MBLG 2071 cdc2 - an example of a CDC mutant - elongated cells at restrictive temperature - cells have copied DNA - these cells get to the end of G2 phase, but for some reason cannot get into M phase - they were later found to be essential cells for transition into M phase from G2 phase - cdc2 kinase (Cdk) is essential for transition from G2 phase to M phase - hence, when the gene encoding this protein is mutated, cells arrest in G2 phase - Cdk levels remain fairly constant throughout the cell cycle and are only activated through interaction with the appropriate cyclin at specific stages - groups of Cdks are names according to phase of the cell cycle in which they are active; G1/S Cdk, S-phase Cdk, M-phase Cdk - also have specific names e.g. cdc2 Cyclin dependent kinases play critical roles in regulating progression through the cell cycle (Hartwell and Nurse) Cyclins Tim Hunt worked with sea urchins and discovered a group of proteins whose level varied cyclically during the cell cycle Hunt names these, cyclins Groups of cyclins are names according to the phase of the cell cycle in which their levels are highest; G1/S cyclins, S-phase cyclins, M-phase cyclins cyclins also have specific names e.g. cyclin B Cyclins regulate the activity of the cyclin dependent kinases (Hunt) cyclins and Cdks Cdk inactive cyclin Cdk active Progression through each of the phases of the cell cycle is controlled by pairs of cyclins and cyclin dependent kinases (Cdk) activated Cdk phosphorylates other proteins

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