Cellular Structures: Microtubules, Filaments, and Extracellular Components

Microtubules

  • Definition and Structure:

    • Hollow rods found in all eukaryotic cells.

    • Constructed from globular proteins called tubulins.

    • Each tubulin protein is a dimer, made of two slightly different polypeptides: α\alpha-tubulin and β\beta-tubulin.

  • Growth and Dynamics:

    • Grow in length by adding tubulin dimers.

    • Can be disassembled and their tubulins reused to build microtubules elsewhere in the cell.

    • Have a distinct polarity due to tubulin dimer orientation: one end (the "plus end") can accumulate or release tubulin dimers at a much higher rate, causing significant growth and shrinkage during cellular activities. The "plus end" designation indicates higher "on" and "off" rates, not exclusively addition.

  • Functions:

    • Shape and support the cell.

    • Serve as tracks for organelles equipped with motor proteins.

    • Guide vesicles from the Endoplasmic Reticulum (ER) to the Golgi apparatus and from the Golgi to the plasma membrane.

    • Involved in the separation of chromosomes during cell division.

Centrosomes and Centrioles

  • Centrosomes:

    • The region in animal cells from which microtubules grow, typically located near the nucleus.

    • Microtubules originating from centrosomes act as compression-resisting girders of the cytoskeleton.

  • Centrioles:

    • Located within the centrosome.

    • Consists of a pair of centrioles, each approximately 250 nm250 \, nm (0.25 μm0.25 \, \mu m) in diameter.

    • Each centriole is composed of nine sets of triplet microtubules arranged in a ring.

    • Although they aid in organizing microtubule assembly in animal cells, many other eukaryotic cells lack centrosomes with centrioles and organize microtubules by alternative means.

    • The two centrioles are typically at right angles to each other.

Cilia and Flagella

  • General Characteristics:

    • Cellular extensions containing microtubules, found in some eukaryotic cells.

    • Responsible for the beating of these structures.

    • Bacterial flagella have a completely different structure.

  • Locomotor Appendages:

    • Many unicellular protists use cilia or flagella for propulsion through water.

    • Sperm of animals, algae, and some plants possess flagella.

  • Fluid Movement:

    • In tissue layers, cilia extending from tightly attached cells can move fluid over the tissue surface.

    • Example: Ciliated lining of the trachea sweeps mucus containing debris out of the lungs.

    • Example: Cilia lining the oviducts in women help move an egg toward the uterus.

  • Differences between Motile Cilia and Flagella:

    • Motile Cilia: Usually occur in large numbers per cell, shorter than flagella, have an alternating power and recovery stroke (like oars of a boat), beating at a rate of about 4040 to 6060 strokes per second.

    • Flagella: Usually limited to one or a few per cell, longer than cilia, have an undulating motion (like a fish tail), driving the cell in the same direction as the flagellum's axis.

Non-motile (Primary) Cilia

  • Function:

    • Can act as a signal-receiving "antenna" for the cell.

    • Generally non-motile, with only one per cell.

    • Almost all vertebrate animal cells possess such a primary cilium.

    • Membrane proteins on these cilia transmit molecular signals from the cell's environment to its interior, triggering signaling pathways that can alter cell activities.

    • Cilium-based signaling is crucial for brain function and embryonic development.

Common Structure of Motile Cilia and Flagella

  • Sheathed Microtubules:

    • Each motile cilium or flagellum has a group of microtubules sheathed in an extension of the plasma membrane.

  • "9+2" Arrangement:

    • Nine doublets of microtubules are arranged in a ring with two single microtubules in the center.

    • This pattern is characteristic of nearly all eukaryotic flagella and motile cilia.

    • Non-motile primary cilia have a "9+0" pattern, lacking the central pair of microtubules.

  • Basal Body:

    • The microtubule assembly is anchored in the cell by a basal body.

    • Structurally very similar to a centriole, with microtubule triplets in a "9+0" pattern.

    • In many animals (including humans), the basal body of the fertilizing sperm's flagellum enters the egg and becomes a centriole.

  • Mechanism of Bending:

    • Bending movements are produced by large motor proteins called dyneins, attached along each outer microtubule doublet.

    • Dynein proteins have two "feet" that "walk" along the microtubule of the adjacent doublet, using ATP for energy.

    • One foot maintains contact while the other releases and reattaches.

    • Flexible cross-linking proteins hold the outer doublets and central microtubules together.

    • The coordinated walking movement on one side of the circle causes the microtubules and the entire organelle to bend, rather than sliding past each other.

Microfilaments (Actin Filaments)

  • Definition and Structure:

    • Thin solid rods, also called actin filaments.

    • Built from molecules of actin, a globular protein.

    • A microfilament is a twisted double chain of actin subunits.

    • Can form structural networks when proteins bind along their side, allowing new filaments to branch.

  • Prevalence: Present in all eukaryotic cells.

  • Structural Role (Bearing Tension):

    • Bear tension (pulling forces) in the cytoskeleton, contrasting with the compression-resisting role of microtubules.

    • Cortical microfilaments: A 3D3D network just inside the plasma membrane that supports cell shape, giving the outer cytoplasmic layer (cortex) a semisolid, gel-like consistency.

    • Microvilli: Bundles of microfilaments form the core of microvilli, delicate projections in cells like nutrient-absorbing intestinal cells, which increase surface area.

  • Role in Cell Motility:

    • Involved in muscle cell contraction: thousands of actin filaments interact with thicker myosin filaments, with myosin projections ("heads") walking along actin to shorten the muscle cell.

    • Amoeboid movement: Localized contractions by actin and myosin enable cells like Amoeba and white blood cells to crawl by extending pseudopodia ("false feet").

    • Cytoplasmic streaming: Actin-protein interactions contribute to the circular flow of cytoplasm within plant cells, speeding organelle movement and material distribution.

Intermediate Filaments

  • Naming and Diameter:

    • Named because their diameter is larger than microfilaments but smaller than microtubules.

  • Prevalence: Found only in the cells of some animals, including vertebrates.

  • Structural Role (Bearing Tension):

    • Specialized for bearing tension, similar to microfilaments.

    • A diverse class of cytoskeletal elements, with each type constructed from a specific molecular subunit (e.g., keratins).

    • Unlike microtubules and microfilaments, which have consistent diameter and composition across eukaryotic cells, intermediate filaments are more varied.

  • Permanence:

    • More permanent fixtures of cells compared to microfilaments and microtubules, which are frequently disassembled and reassembled.

    • Their networks often persist even after cells die (e.g., keratin filaments in the outer layer of skin).

  • Functions:

    • Are especially sturdy and play an important role in reinforcing cell shape.

    • Fix the position of certain organelles; for instance, the nucleus is typically held within a cage of intermediate filaments.

    • Form the nuclear lamina, which lines the interior of the nuclear envelope.

    • Generally function as the permanent framework of the entire cell.

Extracellular Components and Connections Between Cells

  • Cells synthesize and secrete materials extracellularly (outside the cell).

  • These extracellular materials and structures are vital for many essential cellular functions.

Cell Walls of Plants

  • Distinguishing Feature: An extracellular structure that differentiates plant cells from animal cells.

  • Functions:

    • Protects the plant cell.

    • Maintains cell shape.

    • Prevents excessive uptake of water.

    • Provides structural support to the whole plant against gravity.

  • Occurrence: Also found in prokaryotes, some protists, and fungi.

  • Dimensions and Composition:

    • Much thicker than the plasma membrane, ranging from 0.1 μm0.1 \, \mu m to several micrometers.

    • Chemical composition varies by species and cell type, but basic design is consistent.

    • Cellulose microfibrils (a polysaccharide) are synthesized by cellulose synthase and secreted to the extracellular space.

    • Microfibrils are embedded in a matrix of other polysaccharides and proteins, forming a strong fiber-in-ground-substance architectural design (similar to steel-reinforced concrete or fiberglass).

  • Layers:

    • Primary Cell Wall: A relatively thin and flexible wall secreted first by young plant cells.

    • Middle Lamella: A thin, sticky layer rich in pectins (polysaccharides) located between the primary walls of adjacent cells, serving to glue them together.

    • Secondary Cell Wall: Added by some plant cells between the plasma membrane and the primary wall after maturation and growth cessation.

      • Often deposited in several laminated layers.

      • Has a strong and durable matrix, providing significant protection and support (e.g., wood consists mainly of secondary walls).

Extracellular Matrix (ECM) of Animal Cells

  • Lack of Cell Walls: Animal cells do not have cell walls like plant cells.

  • Composition:

    • An elaborate matrix outside the cells.

    • Main ingredients are glycoproteins (proteins with covalently bonded carbohydrates) and other carbohydrate-containing molecules secreted by cells.

  • Key Components:

    • Collagen: The most abundant glycoprotein in the ECM of most animal cells, forming strong fibers outside the cells.

      • Accounts for approximately 40%40\% of the total protein in the human body.

    • Proteoglycans: Collagen fibers are embedded in a network woven out of proteoglycans, which are also secreted by the cells.