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What does the cytoskeleton help provide structure for?
The interior of a eukaryotic cell
What is the cytoskeleton?
A complex network of interconnected filaments and tubules that extends through the cytosol
What does the cytoskeleton provide for cellular function?
Provides architectural framework
What are the characteristics of the cytoskeleton?
High internal organization, dense, fibrous, dynamic, changeable
What does the cytoskeleton enable?
Assumption and maintenance of complex shapes that otherwise wouldn’t be possible
What does the cytoskeleton play a role in?
Cell movement and division, movement of organelles in cytosol, cell signaling, cell-cell adhesion
What is a fundamental feature of the cytoskeleton in both eukaryotes and prokaryotes?
Modularity
What is modularity in terms of the cytoskeleton?
A small number of cytoskeletal elements are deployed in different locations and arranged in different ways to meet the needs of a particular cellular structure
What are the three main structural elements of the cytoskeleton in eukaryotes?
Microtubules, microfilaments, and intermediate filaments
What was the technique of immunofluorescence important for?
Localizing specific proteins to the cytoskeleton
What is each structural element of the cytoskeleton formed by?
The polymerization of a different kind of protein subunit
What is the smallest of the structural elements of the cytoskeleton?
Microfilaments
What is the largest of the sturctural elements of the cytoskeleton?
Microtubules
What is the structure of microtubules?
Hollow tube with a wall consisting of typically 13 protofilaments
What is the structure of microfilaments?
Two intertwined chains of F-actin
What is the structure of intermediate filaments?
Eight protofilaments joined end to end with staggered overlaps
What are septins sometimes called?
The fourth cytoskeleton
What are septins closely associated with?
The contractile ring
What is the contractile ring?
A structure involved in the pinching off of daughter cells during cell division
How do the polymer systems of bacteria and archaea function?
Similarly to microfilaments, microtubules, and intermediate filaments
What is the cytoskeleton best known for its roles in?
Cell motility
What are microfilaments essential components of?
Muscle fibrils
What are microtubules the structural elements of?
Cilia and flagella
Where most cells dynamically regulate, what happens to specific cytoskeletal structures?
They are assembled and disassembled
What do modern cell biologists use to study the cytoskeleton?
Fluorescence microscopy, digital video microscopy, and electron microscopy
What two general groups can microtubules be classified into?
Cytosolic and axonemal
What do cytosolic MTs do in animals?
Maintain axons
What are axons?
Specialized extension of nerve cells
What do cytosolic MTs do in plant cells?
Govern the orientation of cellulose microfibrils
What do cytosolic MTs form that are essential for the movement of chromosomes?
Mitotic and meiotic spindles
What are axonemal microtubules?
Microtubules present in highly ordered bundles in the axonemes of eukaryotic cilia and flagella
What is an axoneme?
The central shaft of a cilium or flagellum consisting of a highly ordered bundle of axonemal MTs and associated proteins
What are the protein building blocks of microtubules?
Tubulin heterodimers
What is the structure of MTs?
Straight, hollow cylinder with an outer diameter 25 nm and inner diameter of 15 nm
What does the MT wall consist of?
Longitudinal arrays of protofilaments
What are protofilaments?
Linear polymer of tubulin subunits
What are protofilaments usually arranged side by side around?
The lumen of the MT
How do the structures of alpha and beta tubulins compare?
They are nearly identical 3D structures despite sharing only 40% amino acid sequence identity
What does each a and b tubulin have?
A binding domain at the N-terminus, a domain in the middle to which colchicine can bind, and a domain at the C-terminus that interacts with MAPs
What does the uniform orientation of tubulin dimers mean?
One end of the protofilament differs structurally from the other
What are tubulin isoforms?
Several closely related but nonidentical genes for each of the a- and b-tubulin subunits
What can some axonemal MTs contain?
Doublet or triplet MTs
What do doublets and triplets contain?
One complete 13-protofilament microtubules and one or two incomplete 10 or 11-protofilament tubules
Where are doublets found?
Cilia and flagella
Where are triplets found?
Basal bodies and centrioles
What are doublets and triplets stabilized by?
Microtubule inner proteins
What do microtubules form by?
The reversible polymerization of tubulin dimers
What do oligomers serve as?
Seeds from which new microtubules can grow in a process called nucleation
What is elongation?
Addition of subunits at either end of an MT after it has been nucleated
What is the lag phase of MT assembly?
The initial slow period of microtubule formation
What is the plateau phase of MT assembly?
When MT assembly is balanced by disassembly
What is critical concentration?
The tubulin heterodimer concentration at which MT assembly is exactly balanced with disassembly
Where does addition of tubulin dimers occur more quickly?
The plus ends of microtubules
What are the two ways that actin exists?
G-actin (monomer) or F-actin (polymer)
What end of the microfilament grows faster?
Barbed (+) end
What do actin-binding proteins do?
Regulate capping, severing, cross-linking, and nucleation; control actin filament organization and turnover
What coordinate polymerization sites?
Signaling pathways
What are the functions of the actin cytoskeleton?
Cell motility, shape, division, intracellular transport
What are two kinds of capping proteins?
CapZ and tropomodulins
What does filamin do?
Bind filaments together at their crossing points and causes microfilaments to become large 3D networks
What does gelsolin do?
Severs microfilaments then caps the newly formed ends
What does ⍺-actinin do?
Makes adhesive connections between filaments in focal contacts and focal adhesions
What does the bundling protein fascin do?
Holds actin in the core of filopodia
What are the actin binding proteins in the microvilli?
Fimbrin and villin
What is the terminal web?
Dense mesh of proteins spectrin and myosin
What are anchored to the terminal web?
Actin filaments
What do membrane linking proteins do?
Connect microfilaments to the membrane of a cell for structural support
What is the formation of dendritic filament networks controlled by?
WASP and the Arp2/3 complex
What is the Arp2/3 complex?
Complex of proteins necessary for branching of microfilaments
What is the WASP?
Complex of proteins that activate the Arp2/3 complex
What do phospholipids and GTPases do?
Regulate the breakdown, formation, and stability of microfilaments
When are Rho family GTPases active?
When bound to GTP
What is myosin?
A family of protein-motors located on the actin cytoskeleton
How many known members does the myosin family have?
At least 24
What are the three main sections of the structure of myosin?
Head domain, neck domain, and tail domain
What is the head domain also known as?
The motor domain
What is the head domain made up of?
A globular protein with active sites to bind actin and ATP
What is the function of the head domain?
To bind to actin and ATP
What does ATP being hydrolyzed cause?
A conformational change that causes the myosin to pull on the actin filaments
What stabilize the head structure?
Light chains
What is the function of light chains?
Allowing the head to move more freely to regulate ATP hydrolysis by the myosin head
What can the tail be bound to?
Cargo or other myosin filaments
What dictate the destination of the myosin?
Specific binding partners in the tail
What is the function of myosin as a whole?
Generation of movement or tension using ATP to exert force on actin filaments
How much efficiency do myosin work with when pulling a moderate load?
50%
What is Myosin I utilized for?
Intracellular transport and membrane interactions
What is Myosin II utilized for?
Muscle contraction and cytokinesis
What is Myosin V utilized for?
Intracellular transport of vesicles and mRNA