Julianne Zedalis; John Eggebrecht

Learning Objectives

In this section, you will explore the following questions:

How do the various components of the cytoskeleton perform their functions?

Connection for AP^® Courses

All cells, from simple bacteria to complex eukaryotes, possess a cytoskeleton composed of different types of protein elements, including microfilaments, intermediate filaments, and microtubules. The cytoskeleton serves a variety of purposes: provides rigidity and shape to the cell, facilitates cellular movement, anchors the nucleus and other organelles in place, moves vesicles through the cell, and pulls replicated chromosomes to the poles of a dividing cell. These protein elements are also integral to the movement of centrioles, flagella, and cilia.

The information presented and the examples highlighted in the section support concepts and Learning Objectives outlined in Big Idea 1 of the AP Biology Curriculum Framework, as shown in the table below.

The Learning Objectives listed in the Curriculum Framework provide a transparent foundation for the AP^® Biology course, an inquiry-based laboratory experience, instructional activities, and AP^® exam questions. A Learning Objective merges required content with one or more of the seven Science Practices.

Big Idea 1	The process of evolution drives the diversity and unity of life.
Enduring Understanding 1.B	Organisms are linked by lines of descent from common ancestry
Essential Knowledge	1.B.1 Organisms share many conserved core processes and features that evolved and are widely distributed among organisms today.
Science Practice	7.2 The student can connect concepts in and across domain(s) to generalize or extrapolate in and/or across enduring understandings and/or big ideas.
Learning Objective	1.15 The student is able to describe specific examples of conserved core biological processes and features shared by all domains or within one domain of life and how these shared, conserved core processes and features support the concept of common ancestry for all organisms.

Teacher Support

Describe the cytoskeleton both as a “skeleton” because it provides the cell with shape and as “muscles” because it allows cells to move. The subunits of the cytoskeleton assemble and disassemble constantly, which is hard to imagine. Stress the concept of “dynamic equilibrium.” A vivid animation may illustrate the point better. Another way to demonstrate this is to have a few students stand in a line and then have a “pool” of students stand nearby. Then, on by one, ask students from the pool to exchange places with a student standing in the line. The line itself can grow and shrink by adding or taking away students in the line, respectively.

Both prokaryotic and eukaryotic cells possess cytoskeletons involved in cell division and shape maintenance. Although the molecular structures of cytoskeleton proteins are similar between two types of cells, the actual amino acid sequences of these proteins show very low levels of homology between the cytoskeleton proteins in prokaryotes and eukaryotes.

The two systems are not closely related.

Although the cytoskeleton of prokaryotes was discovered in the mid 90’s, the misconception that prokaryotes do not have a cytoskeleton is still widespread.

Microfilaments

If you were to remove all the organelles from a cell, would the plasma membrane and the cytoplasm be the only components left? No. Within the cytoplasm, there would still be ions and organic molecules, plus a network of protein fibers that help maintain the shape of the cell, secure some organelles in specific positions, allow cytoplasm and vesicles to move within the cell, and enable cells within multicellular organisms to move. Collectively, this network of protein fibers is known as the cytoskeleton. There are three types of fibers within the cytoskeleton: microfilaments, intermediate filaments, and microtubules (Figure 4.22). Here, we will examine each.

Figure 4.22 Microfilaments thicken the cortex around the inner edge of a cell; like rubber bands, they resist tension. Microtubules are found in the interior of the cell where they maintain cell shape by resisting compressive forces. Intermediate filaments are found throughout the cell and hold organelles in place.

Of the three types of protein fibers in the cytoskeleton, microfilaments are the narrowest. They function in cellular movement, have a diameter of about 7 nm, and are made of two intertwined strands of a globular protein called actin (Figure 4.23). For this reason, microfilaments are also known as actin filaments.

This illustration shows two actin filaments wound together. Each actin filament is composed of many actin subunits connected together to form a chain.

Figure 4.23 Microfilaments are made of two intertwined strands of actin.

Actin is powered by ATP to assemble its filamentous form, which serves as a track for the movement of a motor protein called myosin. This enables actin to engage in cellular events requiring motion, such as cell division in eukaryotic cells and cytoplasmic streaming, which is the circular movement of the cell cytoplasm in plant cells. Actin and myosin are plentiful in muscle cells. When your actin and myosin filaments slide past each other, your muscles contract.

Microfilaments also provide some rigidity and shape to the cell. They can depolymerize (disassemble) and reform quickly, thus enabling a cell to change its shape and move. White blood cells (your body’s infection-fighting cells) make good use of this ability. They can move to the site of an infection and phagocytize the pathogen.

Link to Learning

To see an example of a white blood cell in action, click here and watch a short time-lapse video of the cell capturing two bacteria. It engulfs one and then moves on to the other.

The Human Immunodeficiency Virus (HIV) infects and kills white blood cells. Over time, what affect does this have on the body’s immune system?

The body’s immune system would not be affected by this.
The body’s immune system would not be able to fight off pathogens like bacteria with fewer white blood cells. This can increase the risk of illness in HIV patients.
The body’s immune system, in order to recoup this loss, will produce more WBC’s.
The body’s immune system will fight the pathogens more vigorously in order to compensate for the fewer white blood cells.

Intermediate Filaments

Intermediate filaments are made of several strands of fibrous proteins that are wound together (Figure 4.24). These elements of the cytoskeleton get their name from the fact that their diameter, 8 to 10 nm, is between those of microfilaments and microtubules.

This illustration shows 10 intermediate filament fibers bundled together.

Figure 4.24 Intermediate filaments consist of several intertwined strands of fibrous proteins.

Intermediate filaments have no role in cell movement. Their function is purely structural. They bear tension, thus maintaining the shape of the cell, and anchor the nucleus and other organelles in place. Figure 4.22 shows how intermediate filaments create a supportive scaffolding inside the cell.

The intermediate filaments are the most diverse group of cytoskeletal elements. Several types of fibrous proteins are found in the intermediate filaments. You are probably most familiar with keratin, the fibrous protein that strengthens your hair, nails, and the epidermis of the skin.

Microtubules

As their name implies, microtubules are small hollow tubes. The walls of the microtubule are made of polymerized dimers of α-tubulin and β-tubulin, two globular proteins (Figure 4.25). With a diameter of about 25 nm, microtubules are the widest components of the cytoskeleton. They help the cell resist compression, provide a track along which vesicles move through the cell, and pull replicated chromosomes to opposite ends of a dividing cell. Like microfilaments, microtubules can disassemble and reform quickly.

The left part of this figure is a molecular model of 13 polymerized dimers of alpha- and beta-tubulin joined together to form a hollow tube. The right part of this image shows the tubulin structure as a ring of spheres connected together.

Figure 4.25 Microtubules are hollow. Their walls consist of 13 polymerized dimers of α-tubulin and β-tubulin (right image). The left image shows the molecular structure of the tube.

Microtubules are also the structural elements of flagella, cilia, and centrioles (the latter are the two perpendicular bodies of the centrosome). In fact, in animal cells, the centrosome is the microtubule-organizing center. In eukaryotic cells, flagella and cilia are quite different structurally from their counterparts in prokaryotes, as discussed below.

Flagella and Cilia

To refresh your memory, flagella (singular = flagellum) are long, hair-like structures that extend from the plasma membrane and are used to move an entire cell (for example, sperm, Euglena). When present, the cell has just one flagellum or a few flagella. When cilia (singular = cilium) are present, however, many of them extend along the entire surface of the plasma membrane. They are short, hair-like structures that are used to move entire cells (such as paramecia) or substances along the outer surface of the cell (for example, the cilia of cells lining the Fallopian tubes that move the ovum toward the uterus, or cilia lining the cells of the respiratory tract that trap particulate matter and move it toward your nostrils.)

Despite their differences in length and number, flagella and cilia share a common structural arrangement of microtubules called a “9 + 2 array.” This is an appropriate name because a single flagellum or cilium is made of a ring of nine microtubule doublets, surrounding a single microtubule doublet in the center (Figure 4.26).

This transmission electron micrograph shows a cross section of nine microtubule doublets that form a hollow tube. Another microtubule doublet sits in the center of the tube.

Figure 4.26 This transmission electron micrograph of two flagella shows the 9 + 2 array of microtubules: nine microtubule doublets surround a single microtubule doublet. (credit: modification of work by Dartmouth Electron Microscope Facility, Dartmouth College; scale-bar data from Matt Russell)

Science Practice Connection for AP® Courses

Think About It

The ribosomes in bacterial cells and in human cells are made up of proteins and ribosomal RNA, suggesting that both kinds of cells share a common ancestor cell type. What are examples of other features of cells that provide evidence for common ancestry?

Teacher Support

This question is an application of Learning Objectives 1.15 and Science Practice 7.2 because shared conserved core biological processes and features support the concept of common ancestry for all organisms on Earth.

In both prokaryotes and eukaryotes, DNA is the hereditary material and has the same structure.

The cytosol is made of an aqueous gel. The membrane is a fluid bilayer in which proteins are embedded. Do not use the flagellum as an example. It is convergent evolution.

You have now completed a broad survey of the components of prokaryotic and eukaryotic cells. For a summary of cellular components in prokaryotic and eukaryotic cells, see Table 4.1.

Components of Prokaryotic and Eukaryotic Cells

Cell Component	Function	Present in Prokaryotes?	Present in Animal Cells?	Present in Plant Cells?
Plasma membrane	Separates cell from external environment; controls passage of organic molecules, ions, water, oxygen, and wastes into and out of cell	Yes	Yes	Yes
Cytoplasm	Provides turgor pressure to plant cells as fluid inside the central vacuole; site of many metabolic reactions; medium in which organelles are found	Yes	Yes	Yes
Nucleolus	Darkened area within the nucleus where ribosomal subunits are synthesized.	No	Yes	Yes
Nucleus	Cell organelle that houses DNA and directs synthesis of ribosomes and proteins	No	Yes	Yes
Ribosomes	Protein synthesis	Yes	Yes	Yes
Mitochondria	ATP production/cellular respiration	No	Yes	Yes
Peroxisomes	Oxidizes and thus breaks down fatty acids and amino acids, and detoxifies poisons	No	Yes	Yes
Vesicles and vacuoles	Storage and transport; digestive function in plant cells	No	Yes	Yes
Centrosome	Unspecified role in cell division in animal cells; source of microtubules in animal cells	No	Yes	No
Lysosomes	Digestion of macromolecules; recycling of worn-out organelles	No	Yes	No
Cell wall	Protection, structural support and maintenance of cell shape	Yes, primarily peptidoglycan	No	Yes, primarily cellulose
Chloroplasts	Photosynthesis	No	No	Yes
Endoplasmic reticulum	Modifies proteins and synthesizes lipids	No	Yes	Yes
Golgi apparatus	Modifies, sorts, tags, packages, and distributes lipids and proteins	No	Yes	Yes
Cytoskeleton	Maintains cell’s shape, secures organelles in specific positions, allows cytoplasm and vesicles to move within cell, and enables unicellular organisms to move independently	Yes	Yes	Yes
Flagella	Cellular locomotion	Some	Some	No, except for some plant sperm cells.
Cilia	Cellular locomotion, movement of particles along extracellular surface of plasma membrane, and filtration	Some	Some	No

Table 4.1

4.5 Cytoskeleton