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44 Cards in this Set

  • Front
  • Back

Cellular Movement

• Cell motility involves


– Movement of a cell or organism through the environment


– Movement of the environment past or through the cell – Movement of components in the cell


• Contractility, used to describe shortening of muscle cells, is a specialized form of contractility

Motile Systems

• To generate movement, MTs and MFs provide a scaffold for motor proteins or mechanoenzymes that produce motion at the molecular level


• This is an active process that requires ATP


– ATP may instigate changes in shape of a protein that mediates movemen

Two eukaryotic motility systems

• 1. Interactions between motor proteins and microtubules


– E.g., fast axonal transport in neurons, or the sliding of MTs in cilia and flagella


• 2. Interactions between actin and members of the myosin motor proteins


– E.g., muscle contraction

Intracellular Microtubule-Based Movement

MTs provide a rigid set of tracks for transport of a variety of organelles and vesicles


Traffic toward the minus ends of MTs is considered "inbound"; toward the plus end is "outbound"

Kinesin and Dynein

Microtubule-associated motor proteins walk along the MTs and provide the force needed for movement


Each motor protein will have a preferred direction of movement

Example: Axonal Transport

Fast axonal transport, involves movement of vesicles and organelles along MTs


Organelles can be observed moving along filaments through axoplasm (cytoplasm of axons) at rates of about 2 μ m/sec

Two proteins responsible for fast axonal transport

Kinesin I is involved in ATP- dependent transport toward the plus ends (away from the centrosome), called anterograde axonal transport


Cytoplasmic dynein moves particles (cargo) in the opposite direction, called retrograde axonal transport

Motor Proteins Move Along Microtubules by Hydrolyzing ATP

• Kinesins consist of three parts


– A globular head region that attaches to MTs


– A coiled helical region


– A light-chain region involved in attaching the kinesis to other proteins or organelles

Kinesin movement along MTs

• Kinesin movement looks like "walking" with the two globular head domains taking turns as the front foot


• Each kinesin molecule exhibits processivity


– It can move long distances along a MT before detaching from it by releasing bound ADP and acquiring a new ATP, so that the cycle repeats

Dyneins Can Be Grouped into Two Major Classes:

• Cytoplasmic dynein moves toward the minus ends of MTs


– It is associated with a protein complex called dynactin, which helps link it to cargo


• Axonemal dyneins include at least four different types

Microtubule Motors and the Endomembrane System

• MT motors are important for dynamically shaping the complicated endomembrane system


– E.g., ER membrane extensions can be moved along MTs


• The vesicles to and from the Golgi complex are carried by MT motors on microtubule tracks

Axoneme: Cilia

Cilia: are about 2–10 μm long and have a diameter of 0.25 μm and occur in large numbers on the surface of ciliated cells


They occur in both unicellular and multicellular eukaryotes


Cilia display an oarlike pattern of beating, generating a force parallel to the cell surface

Axoneme: Flagella

Flagella move cells through a fluid environment


Diameter =0.25 μm, length up to 200 μ m


They are limited to one or a few per cell


Move with a propagated bending motion

Structure of Cilia and Flagella

Ciliaandflagellashareacommon structure, the axoneme


Itisconnectedtoabasalbodyand surrounded by an extension of the cell membrane


Betweentheaxonemeandbasalbody is a transition zone in which the MTs take on the pattern characteristic of the axoneme

The basal body

Thebasalbodylookslikea centriole, with 9 sets of tubular structures around the circumference


Eachsetisatripletwiththree MTs that share common walls

Axonemes

Axonemes have a characteristic 􏰀9+2􏰁 pattern, with 9 outer doublets and 2 MTs in the center, the central pair

Axoneme Accessory proteins

• the A and B tubules share a wall in which tektin is a major component


• Each A tubule has a set of sidearms that project from each of the outer
doublets; these consist of
axonemal dynein

Axonemal dynein

• The dynein arms occur in pairs, one inner and one outer arm


• Less frequently, adjacent doublets are joined by interdoublet links that limit the extent of relative movement of doublets, which are linked to each other by a protein called nexin

Radial spokes

At regular intervals, radial spokes project inward toward the central pair


The spokes are thought to be important in translating the sliding of MTs into the bending of the axoneme along with nexin

Microtubule Sliding Within the Axoneme Causes Cilia and Flagella to Bend

• The sliding-microtubule model: sliding of MTs relative to each other is converted into localized bending because the doublets are connected to the central pair and to each other

Dynein Arms Are Responsible for Sliding

The driving force for MT sliding is provided by ATP hydrolysis that is catalyzed by the dynein arms


dynein is responsible for the MT sliding; the dynein arm attaches to and detaches from the B tubule cyclically


Axonemal dynein has multiple subunits, the largest three having ATPase activity

Crosslinks and Spokes Are Responsible for Bending

To bend the doubles must somehow be restrained so that there is resistance to sliding but there is deformation instead


Resistance in bending is provided by:


– the radial spokes that connect the doublets to the central pair


– nexin crosslinks between doublets

Comparing motions of cilia and flagella

The wave-like motion of the flagellum of a sperm cell from a tunicate.


– Note that waves of constant amplitude move continuously from the base to the tip of the flagellum.


The beat of a cilium, which resembles the breast stroke in swimming.

Intracellular Actin-Based Cell Movement:

• Two forms of movement occur as a result of actin function:


1. Cell contraction


2. Cell crawling

Actin Molecular motors: Myosins

Myosins are ATP-dependent motors that exert force on actin filaments


Currently there are 24 known classes of myosins


All have at least one polypeptide chain called the heavy chain, with a globular head group attached to a tail of varying length

The myosin light chains

Myosins typically contain small polypeptides bound to the head group


These light chains play a role in regulating the ATPase


Some myosins bind actin in the tail region, others bind membranes

Myosin functions

• Myosins function in a wide range of cellular events, including


– Muscle contraction
– Cell movement
– Phagocytosis
– Vesicle transport


– Maintainingauditorystructures in humans

Type II myosins

Type II myosins have two heavy chains (each with a globular head, a hinge region, a rodlike tail) and four light chains


They use ATP hydrolysis to cause actin filaments to slide past myosin molecules, resulting in contraction of a cell or group of cells

Many Myosins Move Along Actin Filaments in Short Steps

Myosin II is an efficient motor that 􏰀walks􏰁 along actin


The two heads walk along a protein filament, with the use ATP hydrolysis to change their shape

Structure of skeletal muscle cells

Each muscle fiber contains numerous myofibrils, each of which is divided along its length into repeating units called sarcomeres


Each sarcomere contains bundles of thin filaments (containing actin, troponin and tropomyosin) and thick filaments (containing myosin)

Thick Filaments

Each thick filament consists of hundreds of molecules of myosin, oriented in opposite directions in the two halves of the filament


The myosin is arranged in staggered fashion


Protruding heads of myosin molecules contact the adjacent thin filaments, forming cross-bridges

Thin Filaments

• Thin filaments contain three proteins: F-actin, intertwined with tropomyosin and troponin

The Sliding-Filament Model Explains Muscle Contraction

• According to the model, muscle contraction is due to thin filaments sliding past thick filaments, with no change in length of either

Muscle contraction

• Muscle contraction is the net result of a set of repeated events involving the myosin head


It binds to actin subunits on the thin filament


It undergoes an energy-requiring change in shape


that pulls the thin filament


Then it breaks the association with the actin filament and associates with a site farther along the filament closer to the Z line

The contraction cycle

see picture on pg12

Actin-Based Motility in Non-muscle Cells

• Cell Migration via Lamellipodia i.e. cell crawling, Involves Cycles of:


1. Protrusion of the cells leading edge 2. Attachment to a substrate
3. Translocation
4. Detachment

Extending Protrusions

To crawl, cells extend protrusions at their front, or


leading edge


A thin sheet of cytoplasm is a lamellipodium and a thin-pointed protrusion is a filopodium

Retrograde flow

• Retrograde flow results from actin assembly at the growing tip of the protrusion and rearward translocation of filaments toward the base

Cell Attachment

Attachment, or adhesion, of the cell to the substrate is necessary for cell movement


New sites of attachment are formed at the front of the cell, and contacts at the rear must be broken


Attachment sites are complex structures involving attachment of transmembrane proteins to proteins inside and outside of the cell

Integrins – transmembrane proteins needed for attachment

Integrins on the outside of cells attach to extracellular matrix proteins


Inside the cell integrins are connected to actin filaments via linker proteins


The integrin-dependent attachments are called focal contacts

Translocation and Detachment

Contraction at the rear of the cell squeezes the cell body forward and releases the attachments at the rear


Contraction, due to actin-myosin interactions, is under control of Rho, which activates non-muscle myosin II at the rear of the cell


For movement to occur, new attachments must be balanced by loss of old ones

Chemotaxis

Directional migration occurs through the formation of protrusions predominantly on one side of a cell


Diffusible molecules can act as cues for directional migration; when a cell moves in response to a chemical gradient, it is called chemotaxis


– Chemoattractants: cells move toward a higher concentration of the diffusible molecules


– Chemorepellants: cells move toward a lower concentration of the diffusible molecules

Amoeboid Movement

• Amoebas and white blood cells exhibit a type of crawling called amoeboid movement, which is accompanied by protrusions of pseudopodia


• Involves Cycles of Gelation and Solation of the Actin Cytoskeleton


– Gelation: as a pseudopodium is extended, more material streams forward and congeals at the tip


– Solation: at the rear of the cell, cytosol changes to a more fluid state and streams forward

Movement of Components Within the Cytoplasm of Cells

Cytoplasmic streaming: an actomyosin- dependent movement of cytoplasm in the cell


In plants the process is called cyclosis; a dense set of aligned microfilaments is found near sites where cyclosis occurs