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Advanced Cell Biology
2014 1nd Semester
Department of Animal Science
Chungbuk National University
7th Lecture
1st week : Introduction
3rd week :Research Strategies For Cell Biology
5nd week : Nucleus, Transcription and Splicing
7nd week : Membrane and Channel
9nd week : Membrane Trafficking
11nd week : Cell Signaling
13nd week : Cytoskeleton
15nd week : Cell Cycle
Cytoskeleton : Scaffolds in the Cells
Architectural scaffolds : Supports constructions
Three major cytoskeleton components
Microfilaments Microtubule Intermediate filaments
(Actin) (Tubulin) (Keratin, vimentin..)
Microfilaments
Cytoskeletons as “Road” in the Cell
Highway “고속도로”
Interstate Highway = Microtubule?
“Local Roads”
Microfilaments
(Actin)
Actin
Monomeric Actin (42kDa)
(G-Actin)
Filamentous Actin
(F-Actin)
Actin is the second abundant protein in the world
Polymerization
Depolymerization
Actin Filament:
the main ‘building block’ of cytoskeleton
Borish & Svitskina
Roles of Actin Filament Remodeling in various cellular process
CytokinesisCell Motility
PhagocytosisVesicle Transport
Asymmetric spindle Positioning in Oocyte
Actin Filament is dynamic
Formation of Actin filaments
• When?
• How long?
• Where?
• How looks like?
New Filament Generations (Actin Nucleators:Arp2/3, Formins, Spire)
Capping (CapZ, Tropomodulin)
Depolymerizations/Severing (Cofillin, Gelsolin)
Monomeric Actin Binding Protein (Profilin, Thymosin beta-4)
Actin Nucleators Determine type of Filament Structure
Arp2/3 complex
:59 Annual Reviews AR131-19
REGULATION OF ACTIN FILAMENT NETWORK 651
ure 1 Electron micrographs of branched actin filaments. (a) Electron micrograph of
leading edge of a migrating Xenopus keratocyte prepared by detergent extraction and
ry shadowing. (b) Detail of boxed region from (a), with a branched filament network
ed in blue. (c) Gold-labeled antibody to ARPC2 localized to a branch point in the
ment network at the leading edge. (d ) Branch formed in vitro by pure actin filaments
Acanthamoeba Arp2/3 complex. (a–c) modified from 14. (d ) Modified from 13.
p2/3 complex (hereafter referred to as nucleation promoting factors), are being
Formins
Straight actin bundles
In Stress fibers and
filopodia
Branched actin filaments
in Lamellipodia
Spire
Straight Fiber
Oocyte development
Vesicle Transport
Arp2/3 complex:
6 Protein complex makes branched networks
Arp2
Arp3
ARPC3ARPC1
ARPC5
ARPC4
ARPC2
Dominguez and Namgoong, Comphrensive Biophysics 2012
Arp2
Arp3
ARPC3ARPC1
ARPC5
ARPC4
ARPC2mother filament
barbed end
barbedend
Nucleation Promoting Factor (NPF)
18 85 161 200 245 436 498
N-WASP Pro-rich
26 136 200 270 400 501
WH1 GBDB AW CW
WAVE2 Pro-richhelix B AW C
JMY MA spectrin-like AW CW WPro-rich
40 300 620 790 855 983
profilin-actinregulation/localization
Prototypical NPF AW CWWPro-rich
actin Arp2/3
167 202 290 303 360 465
WASH
Pro-richWHD1 AW CWHD2
Arp2/3 complex WCA + actin transition complex activated complex
+
branch
70º
2
4
5
1
Arp2
Arp3
3
Nucleation Promoting Factor (NPF) are essential for
Arp2/3 mediated filament nucleations
Formins
Rho actin
Pro-rich (FH1) FH2CCDD DADDIDGBD
73 130 377 452 570 736 1175 1200
autoinhibition
profilin-actin
DD
DID
DAD
Complex of DID and DAD
DID
GTP-Rho
DD
GBD
Binding of Rho GTPase
linker
lasso
knobcoiled coil
post
FH2 Domain
3
1
4
21
2 4
barbed end
barbed end
3
1
4
2
1
2 4
3
• Responsible for the straight bundles of filaments (Stress fiber,
Fillopodia)
• 15 formins (8 subclass) in human and mouse
• Caps fast growing ends (barbed end) of actin filament and enhance
elongation rates
• Regulated by autoinhibition and small GTPase (Rho)
Signaling activating Actin nucleators
Rho-family GTPases (Rac, RhoA, cdc42) play crucial roles in activation of Actin Nucleators
Rac -> WAVE -> Arp2/3
Cdc42 -> N-WASP -> Arp2/3
RhoA -> Formin
1253
1128
158 445 507
Sra1
Nap1
90 172 202 498 559
W C Ahelix
meander
Pro-rich region
WAVE1
Abi2
75
HSPC300
L697
Y704
L841
F844
W845E434 F626 M632R190C179
basic
Rac1-binding
SH3
Nap1
HSPC300
Sra1
Abi2
WAVE1
WAVE1 Polybasic region
(proposed membrane-binding site)
proposed Rac1-binding site
W
C
C-terminal part polybasic region and Pro-rich sequence of WAVE1
substituted by a short linker (disordered in structure)
Pro-rich region and SH3 domain
removed for crystallization
disordered
Actin Depolymerization
- ADF/Cofillin : Bind to Actin Filaments (ADP-Actin)
- Favor Depolymerization of Actin
- ADF/Cofillin signaling
RhoA->ROCK-> LIM Kinase -> Cofillin Phosphorylation (Inactivation)
Slingshot Phosphatase
Dephosphorylated Cofilin (Active)
Actin Capping Proteins
- Barbed End (Fast-Growing End) Capper
CapZ : Stop Elongation of Actin Filaments
- Pointed End Capper
Tropomodulin : Protect Actin filaments from Depolymerization
Capping Proteins : inhibit elongation and stabilize
actin filaments
CapZαβ : Capping fast growing (Barbed) end
Gelsolin : Severing and Capping Proteins
Tropomyosin + Tropomodulin : Guardian of actin filament
Tropomodulin
Tropomyosin
Dendritic Nucleation Model
Myosin : Molecular Motor on Microfilaments
Cell bio7
Myosin – Actin in Skeletal Muscle
Cell bio7
Myosin walk on actin filaments
- Except Myosin VI, every myosin move from
Slow growing end (pointed end)
- Hydrolysis of ATP provides energy for the movement
Regulation of Myosin Activities
Skeletal Muscle : Calcium mediated
Troponin-Tropomyosin
Smooth Muscle :
Myosin Light Chain Kinase
Myosin Light Chain Phosphatase
Cell Motility
Actin Polymerization is the main driving force for cell migration
Actin filaments in Lamellipodia is mainly formed as branched filaments by
Arp2/3 complex
Actin filaments in Fillopodia is mainly formed as straight bundle by Formins
and Arp2/3 complex
Cytokinesis
http://bement.molbio.wisc.edu/node/13
Endocytosis
Asymmetric division
Actin Cap
4N
GV GVBD
Spindle
Formations
Spindle
Migration
MII
Arrest
2N
2N
1N
1N
Fertilization
Polar Body
Adapted from Yi and RI, Cytoskeleton (2012)
Actin Nucleators in Asymmetric Spindle
Migrations
Gene Phenotype Method Referenc
es
Formin
(FMN2)
Failure of Spindle
Migration
Symmetric division
Knockout mouse
fmn2 (-/-)
Spire Failture of Spindle
Migration
Failure in Cytokinesis
siRNA
knockdown
How about Arp2/3 and Nucleation Promoting Factors?
Microtubule
Consists with alpha and beta Tubulin (Binds GTP or GDP)
Alpha and Beta Tubulin
Cell bio7
Cell bio7
Mitotic spindle
Centrosome
Cell bio7
Microtubule-binding proteins
Microtubule-based Motors
- Kinesin : Minus – Plus End Directed Motor Proteins
- Dynein : Plus -> Minus Directed Motor Proteins
Kinesin walking on microtubule
Moving vesicle along with microtubule
Cell bio7

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Cell bio7

  • 1. Advanced Cell Biology 2014 1nd Semester Department of Animal Science Chungbuk National University 7th Lecture
  • 2. 1st week : Introduction 3rd week :Research Strategies For Cell Biology 5nd week : Nucleus, Transcription and Splicing 7nd week : Membrane and Channel 9nd week : Membrane Trafficking 11nd week : Cell Signaling 13nd week : Cytoskeleton 15nd week : Cell Cycle
  • 3. Cytoskeleton : Scaffolds in the Cells Architectural scaffolds : Supports constructions
  • 4. Three major cytoskeleton components Microfilaments Microtubule Intermediate filaments (Actin) (Tubulin) (Keratin, vimentin..) Microfilaments
  • 5. Cytoskeletons as “Road” in the Cell Highway “고속도로” Interstate Highway = Microtubule? “Local Roads” Microfilaments (Actin)
  • 6. Actin Monomeric Actin (42kDa) (G-Actin) Filamentous Actin (F-Actin) Actin is the second abundant protein in the world Polymerization Depolymerization
  • 7. Actin Filament: the main ‘building block’ of cytoskeleton Borish & Svitskina
  • 8. Roles of Actin Filament Remodeling in various cellular process CytokinesisCell Motility PhagocytosisVesicle Transport Asymmetric spindle Positioning in Oocyte
  • 9. Actin Filament is dynamic Formation of Actin filaments • When? • How long? • Where? • How looks like?
  • 10. New Filament Generations (Actin Nucleators:Arp2/3, Formins, Spire) Capping (CapZ, Tropomodulin) Depolymerizations/Severing (Cofillin, Gelsolin) Monomeric Actin Binding Protein (Profilin, Thymosin beta-4)
  • 11. Actin Nucleators Determine type of Filament Structure Arp2/3 complex :59 Annual Reviews AR131-19 REGULATION OF ACTIN FILAMENT NETWORK 651 ure 1 Electron micrographs of branched actin filaments. (a) Electron micrograph of leading edge of a migrating Xenopus keratocyte prepared by detergent extraction and ry shadowing. (b) Detail of boxed region from (a), with a branched filament network ed in blue. (c) Gold-labeled antibody to ARPC2 localized to a branch point in the ment network at the leading edge. (d ) Branch formed in vitro by pure actin filaments Acanthamoeba Arp2/3 complex. (a–c) modified from 14. (d ) Modified from 13. p2/3 complex (hereafter referred to as nucleation promoting factors), are being Formins Straight actin bundles In Stress fibers and filopodia Branched actin filaments in Lamellipodia Spire Straight Fiber Oocyte development Vesicle Transport
  • 12. Arp2/3 complex: 6 Protein complex makes branched networks Arp2 Arp3 ARPC3ARPC1 ARPC5 ARPC4 ARPC2 Dominguez and Namgoong, Comphrensive Biophysics 2012 Arp2 Arp3 ARPC3ARPC1 ARPC5 ARPC4 ARPC2mother filament barbed end barbedend Nucleation Promoting Factor (NPF)
  • 13. 18 85 161 200 245 436 498 N-WASP Pro-rich 26 136 200 270 400 501 WH1 GBDB AW CW WAVE2 Pro-richhelix B AW C JMY MA spectrin-like AW CW WPro-rich 40 300 620 790 855 983 profilin-actinregulation/localization Prototypical NPF AW CWWPro-rich actin Arp2/3 167 202 290 303 360 465 WASH Pro-richWHD1 AW CWHD2 Arp2/3 complex WCA + actin transition complex activated complex + branch 70º 2 4 5 1 Arp2 Arp3 3 Nucleation Promoting Factor (NPF) are essential for Arp2/3 mediated filament nucleations
  • 14. Formins Rho actin Pro-rich (FH1) FH2CCDD DADDIDGBD 73 130 377 452 570 736 1175 1200 autoinhibition profilin-actin DD DID DAD Complex of DID and DAD DID GTP-Rho DD GBD Binding of Rho GTPase linker lasso knobcoiled coil post FH2 Domain 3 1 4 21 2 4 barbed end barbed end 3 1 4 2 1 2 4 3 • Responsible for the straight bundles of filaments (Stress fiber, Fillopodia) • 15 formins (8 subclass) in human and mouse • Caps fast growing ends (barbed end) of actin filament and enhance elongation rates • Regulated by autoinhibition and small GTPase (Rho)
  • 15. Signaling activating Actin nucleators Rho-family GTPases (Rac, RhoA, cdc42) play crucial roles in activation of Actin Nucleators Rac -> WAVE -> Arp2/3 Cdc42 -> N-WASP -> Arp2/3 RhoA -> Formin
  • 16. 1253 1128 158 445 507 Sra1 Nap1 90 172 202 498 559 W C Ahelix meander Pro-rich region WAVE1 Abi2 75 HSPC300 L697 Y704 L841 F844 W845E434 F626 M632R190C179 basic Rac1-binding SH3 Nap1 HSPC300 Sra1 Abi2 WAVE1 WAVE1 Polybasic region (proposed membrane-binding site) proposed Rac1-binding site W C C-terminal part polybasic region and Pro-rich sequence of WAVE1 substituted by a short linker (disordered in structure) Pro-rich region and SH3 domain removed for crystallization disordered
  • 17. Actin Depolymerization - ADF/Cofillin : Bind to Actin Filaments (ADP-Actin) - Favor Depolymerization of Actin - ADF/Cofillin signaling RhoA->ROCK-> LIM Kinase -> Cofillin Phosphorylation (Inactivation) Slingshot Phosphatase Dephosphorylated Cofilin (Active)
  • 18. Actin Capping Proteins - Barbed End (Fast-Growing End) Capper CapZ : Stop Elongation of Actin Filaments - Pointed End Capper Tropomodulin : Protect Actin filaments from Depolymerization
  • 19. Capping Proteins : inhibit elongation and stabilize actin filaments CapZαβ : Capping fast growing (Barbed) end Gelsolin : Severing and Capping Proteins
  • 20. Tropomyosin + Tropomodulin : Guardian of actin filament Tropomodulin Tropomyosin
  • 22. Myosin : Molecular Motor on Microfilaments
  • 24. Myosin – Actin in Skeletal Muscle
  • 26. Myosin walk on actin filaments - Except Myosin VI, every myosin move from Slow growing end (pointed end) - Hydrolysis of ATP provides energy for the movement
  • 27. Regulation of Myosin Activities Skeletal Muscle : Calcium mediated Troponin-Tropomyosin Smooth Muscle : Myosin Light Chain Kinase Myosin Light Chain Phosphatase
  • 28. Cell Motility Actin Polymerization is the main driving force for cell migration
  • 29. Actin filaments in Lamellipodia is mainly formed as branched filaments by Arp2/3 complex Actin filaments in Fillopodia is mainly formed as straight bundle by Formins and Arp2/3 complex
  • 33. Asymmetric division Actin Cap 4N GV GVBD Spindle Formations Spindle Migration MII Arrest 2N 2N 1N 1N Fertilization Polar Body Adapted from Yi and RI, Cytoskeleton (2012)
  • 34. Actin Nucleators in Asymmetric Spindle Migrations Gene Phenotype Method Referenc es Formin (FMN2) Failure of Spindle Migration Symmetric division Knockout mouse fmn2 (-/-) Spire Failture of Spindle Migration Failure in Cytokinesis siRNA knockdown How about Arp2/3 and Nucleation Promoting Factors?
  • 35. Microtubule Consists with alpha and beta Tubulin (Binds GTP or GDP) Alpha and Beta Tubulin
  • 42. Microtubule-based Motors - Kinesin : Minus – Plus End Directed Motor Proteins - Dynein : Plus -> Minus Directed Motor Proteins
  • 43. Kinesin walking on microtubule
  • 44. Moving vesicle along with microtubule