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Learning Objectives

Learning Objectives

In this section, you will explore the following questions

  • What is the relationship between the structure and function of the components of the endomembrane system, especially with regard to the synthesis of proteins?

Connection for AP® Courses

Connection for AP® Courses

In addition to the presence of nuclei, eukaryotic cells are distinguished by an endomembrane system that includes the plasma membrane, nuclear envelope, lysosomes, vesicles, endoplasmic reticulum, and Golgi apparatus. These subcellular components work together to modify, tag, package, and transport proteins and lipids. The rough endoplasmic reticulum (RER) with its attached ribosomes is the site of protein synthesis and modification. The smooth endoplasmic reticulum (SER) synthesizes carbohydrates, lipids including phospholipids and cholesterol, and steroid hormones; engages in the detoxification of medications and poisons; and stores calcium ions. Lysosomes digest macromolecules, recycle worn-out organelles, and destroy pathogens. Just like your body uses different organs that work together, cells use these organelles together to perform specific functions. For example, proteins that are synthesized in the RER then travel to the Golgi apparatus for modification and packaging for either storage or transport. If these proteins are hydrolytic enzymes, they can be stored in lysosomes. Mitochondria produce the energy needed for these processes. This functional flow through several organelles, a process which is dependent on energy produced by yet another organelle, serves as a hallmark illustration of the cell’s complex, interconnected dependence on its organelles.

Information presented and the examples highlighted in the section support concepts and Learning Objectives outlined in Big Idea 2 and Big Idea 4 of the AP® Biology Curriculum Framework. 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 2 Biological systems utilize free energy and molecular building blocks to grow, to reproduce, and to maintain dynamic homeostasis.
Enduring Understanding 2.B Growth, reproduction, and dynamic homeostasis require that cells create and maintain internal environments that are different from their external environments.
Essential Knowledge 2.B.3 Eukaryotic cells maintain internal membranes that partition the cell into specialized regions.
Science Practice 6.2 The student can construct explanations of phenomena based on evidence produced through scientific practices.
Learning Objective 2.13 The student is able to explain how internal membranes and organelles contribute to cell functions.
Big Idea 4 Biological systems interact, and these systems and their interactions possess complex properties.
Enduring Understanding 4.A Interactions within biological systems lead to complex properties.
Essential Knowledge 4.A.2 The structure and function of subcellular components, and their interactions, provide essential cellular processes.
Learning Objective 4.5 The student is able to construct explanations based on scientific evidence as to how interactions of subcellular structures provide essential functions.

The Science Practices Assessment Ancillary contains additional test questions for this section that will help you prepare for the AP exam. These questions address the following standards:


[APLO 4.6]

The Endoplasmic Reticulum

The Endoplasmic Reticulum

The endomembrane system (endo = within) is a group of membranes and organelles (Figure 4.18) in eukaryotic cells that work together to modify, package, and transport lipids and proteins. It includes the nuclear envelope, lysosomes, and vesicles, which we have already mentioned, and the endoplasmic reticulum and Golgi apparatus, which we will cover shortly. Although not technically within the cell, the plasma membrane is included in the endomembrane system because, as you will see, it interacts with the other endomembranous organelles. The endomembrane system does not include the membranes of either mitochondria or chloroplasts.

Visual Connection

The left part of this figure shows the rough ER with an integral membrane protein embedded in it. The part of the protein facing the inside of the ER has a carbohydrate attached to it. The protein is shown leaving the ER in a vesicle that fuses with the cis side of the Golgi apparatus. The Golgi apparatus consists of several layers of membranes, called cisternae. As the protein passes through the cisternae, it is further modified by the addition of more carbohydrates. Eventually, it leaves the trans face of the Golgi in a vesicle. The vesicle fuses with the cell membrane so that the carbohydrate that was on the inside of the vesicle now faces the outside of the membrane. At the same time, the contents of the vesicle are ejected from the cell.
Figure 4.18 Membrane and secretory proteins are synthesized in the rough endoplasmic reticulum (RER). The RER also sometimes modifies proteins. In this illustration, a (green) integral membrane protein in the ER is modified by attachment of a (purple) carbohydrate. Vesicles with the integral protein buds from the ER and fuse with the cis face of the Golgi apparatus. As the protein passes along the Golgi’s cisternae, it is further modified by the addition of more carbohydrates. After its synthesis is complete, it exits as an integral membrane protein of the vesicle that buds from the Golgi’s trans face, and when the vesicle fuses with the cell membrane, the protein becomes an integral portion of that cell membrane. (credit: modification of work by Magnus Manske)

If a peripheral membrane protein were synthesized inside the lumen of the ER, would it end up on the inside or outside of the plasma membrane?

  1. The vesicle travels from the endoplasmic reticulum to get embedded in the plasma membrane.
  2. The vesicle travels from the Golgi apparatus to the plasma membrane to release the protein outside.
  3. The vesicle travels from the endoplasmic reticulum to the plasma membrane, and returns to the Golgi apparatus to get modified.
  4. The vesicle moves from the endoplasmic reticulum into the cytoplasmic area, remaining there.

The endoplasmic reticulum (ER) (Figure 4.18) is a series of interconnected membranous sacs and tubules that collectively modifies proteins and synthesizes lipids. However, these two functions are performed in separate areas of the ER: the rough ER and the smooth ER, respectively.

The hollow portion of the ER tubules is called the lumen or cisternal space. The membrane of the ER, which is a phospholipid bilayer embedded with proteins, is continuous with the nuclear envelope.

Rough ER

The rough endoplasmic reticulum (RER) is so named because the ribosomes attached to its cytoplasmic surface give it a studded appearance when viewed through an electron microscope (Figure 4.19).

In this transmission electron micrograph, the nucleus is the most prominent feature. The nucleolus is a circular, dark region inside the nucleus. A nuclear pore can be seen in the nuclear envelope that surrounds the nucleus. The rough endoplasmic reticulum surrounds the nucleus, appearing as many layers of membranes. A mitochondrion sits between the layers of the ER membrane.
Figure 4.19 This transmission electron micrograph shows the rough endoplasmic reticulum and other organelles in a pancreatic cell. (credit: modification of work by Louisa Howard)

Ribosomes transfer their newly synthesized proteins into the lumen of the RER where they undergo structural modifications, such as folding or the acquisition of side chains. These modified proteins will be incorporated into cellular membranes—the membrane of the ER or those of other organelles—or secreted from the cell (such as protein hormones; enzymes). The RER also makes phospholipids for cellular membranes.

If the phospholipids or modified proteins are not destined to stay in the RER, they will reach their destinations via transport vesicles that bud from the RER’s membrane (Figure 4.18).

Since the RER is engaged in modifying proteins (such as enzymes, for example) that will be secreted from the cell, you would be correct in assuming that the RER is abundant in cells that secrete proteins. This is the case with cells of the liver, for example.

Smooth ER

The smooth endoplasmic reticulum (SER) is continuous with the RER but has few or no ribosomes on its cytoplasmic surface (Figure 4.18). Functions of the SER include synthesis of carbohydrates, lipids, and steroid hormones; detoxification of medications and poisons; and storage of calcium ions.

In muscle cells, a specialized SER called the sarcoplasmic reticulum is responsible for storage of the calcium ions that are needed to trigger the coordinated contractions of the muscle cells.

Link to Learning

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You can watch an excellent animation of the endomembrane system here. At the end of the animation, there is a short self-assessment.

Explain how the nucleus and the endomembrane system work together for protein synthesis.

  1. The endomembrane system processes and ships proteins specified by the nucleus. In the nucleus, DNA is used to make RNA, which exits the nucleus and enters the cytoplasm of the cell. The ribosomes on the rough ER use the RNA to create the different types of protein needed by the body.
  2. The endomembrane system processes and ships proteins specified by the nucleus. From the nucleus, RNA exits and enters the cytoplasm of the cell. The ribosomes on the rough ER use the RNA to create the different types of protein needed by the body.
  3. The endomembrane system processes and ships proteins specified by the nucleus. In the nucleus, DNA is used to make RNA, which exits the nucleus and enters the cytoplasm of the cell. The smooth ER uses the RNA to create the different types of protein needed by the body.
  4. The endomembrane system processes and ships proteins specified by the nucleus. In the nucleus, DNA is used to make RNA, which exits the nucleus and enters the cytoplasm of the cell. The ribosomes on the smooth ER use the RNA to create the different types of protein needed by the body.

Career Connection

Cardiologist

Heart disease is the leading cause of death in the United States. This is primarily due to our sedentary lifestyle and our high trans-fat diets.

Heart failure is just one of many disabling heart conditions. Heart failure does not mean that the heart has stopped working. Rather, it means that the heart can not pump with sufficient force to transport oxygenated blood to all the vital organs. Left untreated, heart failure can lead to kidney failure and failure of other organs.

The wall of the heart is composed of cardiac muscle tissue. Heart failure occurs when the endoplasmic reticula of cardiac muscle cells do not function properly. As a result, an insufficient number of calcium ions are available to trigger a sufficient contractile force.

Cardiologists (cardi- = heart; -ologist = one who studies) are doctors who specialize in treating heart diseases, including heart failure. Cardiologists can make a diagnosis of heart failure via physical examination, results from an electrocardiogram (ECG—a test that measures the electrical activity of the heart), a chest X-ray to see whether the heart is enlarged, and other tests. If heart failure is diagnosed, the cardiologist will typically prescribe appropriate medications and recommend a reduction in table salt intake and a supervised exercise program.

The Golgi Apparatus

The Golgi Apparatus

We have already mentioned that vesicles can bud from the ER and transport their contents elsewhere, but where do the vesicles go? Before reaching their final destination, the lipids or proteins within the transport vesicles still need to be sorted, packaged, and tagged so that they wind up in the right place. Sorting, tagging, packaging, and distribution of lipids and proteins takes place in the Golgi apparatus (also called the Golgi body): a series of flattened membranes (Figure 4.20).

In this transmission electron micrograph, the Golgi apparatus appears as a stack of membranes surrounded by unnamed organelles.
Figure 4.20 The Golgi apparatus in this white blood cell is visible as a stack of semicircular, flattened rings in the lower portion of the image. Several vesicles can be seen near the Golgi apparatus. (credit: modification of work by Louisa Howard)

The receiving side of the Golgi apparatus is called the cis face. The opposite side is called the trans face. The transport vesicles that formed from the ER travel to the cis face, fuse with it, and empty their contents into the lumen of the Golgi apparatus. As the proteins and lipids travel through the Golgi, they undergo further modifications that allow them to be sorted. The most frequent modification is the addition of short chains of sugar molecules. These newly modified proteins and lipids are then tagged with phosphate groups or other small molecules so that they can be routed to their proper destinations.

Finally, the modified and tagged proteins are packaged into secretory vesicles that bud from the trans face of the Golgi. While some of these vesicles deposit their contents into other parts of the cell where they will be used, other secretory vesicles fuse with the plasma membrane and release their contents outside the cell.

In another example of form following function, cells that engage in a great deal of secretory activity, such as cells of the salivary glands that secrete digestive enzymes or cells of the immune system that secrete antibodies, have an abundance of Golgi.

In plant cells, the Golgi apparatus has the additional role of synthesizing polysaccharides, some of which are incorporated into the cell wall and some of which are used in other parts of the cell.

Career Connection

Geneticist

Many diseases arise from genetic mutations that prevent the synthesis of critical proteins. One such disease is Lowe disease (also called oculocerebrorenal syndrome, because it affects the eyes, brain, and kidneys). In Lowe disease, there is a deficiency in an enzyme localized to the Golgi apparatus. Children with Lowe disease are born with cataracts, typically develop kidney disease after the first year of life, and may have impaired mental abilities.

Lowe disease is a genetic disease caused by a mutation on the X chromosome. The X chromosome is one of the two human sex chromosomes, as these chromosomes determine a person's sex. Females possess two X chromosomes, while males possess one X and one Y chromosome, In females, the genes on only one of the two X chromosomes are expressed. Therefore, females who carry the Lowe disease gene on one of their X chromosomes have a 50/50 chance of having the disease. However, males only have one X chromosome and the genes on this chromosome are always expressed. Therefore, males will always have Lowe disease if their X chromosome carries the Lowe disease gene. The location of the mutated gene, as well as the locations of many other mutations that cause genetic diseases, has now been identified. Through prenatal testing, a woman can find out if the fetus she is carrying may be afflicted with one of several genetic diseases.

Geneticists analyze the results of prenatal genetic tests and may counsel pregnant women on available options. They may also conduct genetic research that leads to new drugs or foods, or perform DNA analyses that are used in forensic investigations.

Lysosomes

Lysosomes

In addition to their role as the digestive component and organelle-recycling facility of animal cells, lysosomes are considered to be parts of the endomembrane system. Lysosomes also use their hydrolytic enzymes to destroy pathogens (disease-causing organisms) that might enter the cell. A good example of this occurs in a group of white blood cells called macrophages, which are part of your body’s immune system. In a process known as phagocytosis or endocytosis, a section of the plasma membrane of the macrophage invaginates (folds in) and engulfs a pathogen. The invaginated section, with the pathogen inside, then pinches itself off from the plasma membrane and becomes a vesicle. The vesicle fuses with a lysosome. The lysosome’s hydrolytic enzymes then destroy the pathogen (Figure 4.21).

In this illustration, a macrophage consuming a bacterium is shown. As the bacterium is consumed by endocytosis, it is encapsulated in a vesicle. A lysosome fuses with the vesicle and digestive enzymes inside the lysosome digest the bacterium. The undigested material is expelled from the macrophage through the process of exocytosis.
Figure 4.21 A macrophage has engulfed (phagocytized) a potentially pathogenic bacterium, which then fuses with a lysosome within the cell to destroy the pathogen. Other organelles are present in the cell but for simplicity are not shown.

Science Practice Connection for AP® Courses

Activity

Homemade Cell Project. Using inexpensive and common household items, create a model of a specific eukaryotic cell (e.g., neuron, white blood cell, plant root cell, or Paramecium) that demonstrates how at least three organelles work together to perform a specific function.

Think About It

A certain cell type functions primarily to synthesize proteins for export. What is the most likely route the newly made protein takes through the cell? Justify your prediction.