Eukaryotic Cells

Learning Objectives

By the end of this section, you will be able to do the following:

  • Describe the structure of eukaryotic cells
  • Compare animal cells with plant cells
  • State the role of the plasma membrane
  • Summarize the functions of the major cell organelles

Have you ever heard the phrase “form follows function?” It’s a philosophy that many industries follow. In architecture, this means that buildings should be constructed to support the activities that will be carried out inside them. For example, a skyscraper should include several elevator banks. A hospital should have its emergency room easily accessible.

Our natural world also utilizes the principle of form following function, especially in cell biology, and this will become clear as we explore eukaryotic cells ( (Figure)). Unlike prokaryotic cells, eukaryotic cells have: 1) a membrane-bound nucleus; 2) numerous membrane-bound organelles such as the endoplasmic reticulum, Golgi apparatus, chloroplasts, mitochondria, and others; and 3) several, rod-shaped chromosomes. Because a membrane surrounds eukaryotic cell’s nucleus, it has a “true nucleus.” The word “organelle” means “little organ,” and, as we already mentioned, organelles have specialized cellular functions, just as your body’s organs have specialized functions.

At this point, it should be clear to you that eukaryotic cells have a more complex structure than prokaryotic cells. Organelles allow different functions to be compartmentalized in different areas of the cell. Before turning to organelles, let’s first examine two important components of the cell: the plasma membrane and the cytoplasm.

Art Connection

These figures show the major organelles and other cell components of (a) a typical animal cell and (b) a typical eukaryotic plant cell. The plant cell has a cell wall, chloroplasts, plastids, and a central vacuole—structures not in animal cells. Most cells do not have lysosomes or centrosomes.

Part a: This illustration shows a typical eukaryotic animal cell, which is egg shaped. The fluid inside the cell is called the cytoplasm, and the cell is surrounded by a cell membrane. The nucleus takes up about one-half the width of the cell. Inside the nucleus is the chromatin, which is composed of DNA and associated proteins. A region of the chromatin is condensed into the nucleolus, a structure where ribosomes are synthesized. The nucleus is encased in a nuclear envelope, which is perforated by protein-lined pores that allow entry of material into the nucleus. The nucleus is surrounded by the rough and smooth endoplasmic reticulum, or ER. The smooth ER is the site of lipid synthesis. The rough ER has embedded ribosomes that give it a bumpy appearance. It synthesizes membrane and secretory proteins. In addition to the ER, many other organelles float inside the cytoplasm. These include the Golgi apparatus, which modifies proteins and lipids synthesized in the ER. The Golgi apparatus is made of layers of flat membranes. Mitochondria, which produce food for the cell, have an outer membrane and a highly folded inner membrane. Other, smaller organelles include peroxisomes that metabolize waste, lysosomes that digest food, and vacuoles. Ribosomes, responsible for protein synthesis, also float freely in the cytoplasm and are depicted as small dots. The last cellular component shown is the cytoskeleton, which has four different types of components: microfilaments, intermediate filaments, microtubules, and centrosomes. Microfilaments are fibrous proteins that line the cell membrane and make up the cellular cortex. Intermediate filaments are fibrous proteins that hold organelles in place. Microtubules form the mitotic spindle and maintain cell shape. Centrosomes are made of two tubular structures at right angles to one another. They form the microtubule-organizing center.

Part b: This illustration depicts a typical eukaryotic plant cell. The nucleus of a plant cell contains chromatin and a nucleolus, the same as an animal cell. Other structures that the plant cell has in common with the animal cell include rough and smooth endoplasmic reticulum, the Golgi apparatus, mitochondria, peroxisomes, and ribosomes. The fluid inside the plant cell is called the cytoplasm, just as it is in an animal cell. The plant cell has three of the four cytoskeletal components found in animal cells: microtubules, intermediate filaments, and microfilaments. Plant cells do not have centrosomes. Plant cells have four structures not found in animals cells: chloroplasts, plastids, a central vacuole, and a cell wall. Chloroplasts are responsible for photosynthesis; they have an outer membrane, an inner membrane, and stack of membranes inside the inner membrane. The central vacuole is a very large, fluid-filled structure that maintains pressure against the cell wall. Plastids store pigments. The cell wall is outside the cell membrane.

If the nucleolus were not able to carry out its function, what other cellular organelles would be affected?

The Plasma Membrane

Like prokaryotes, eukaryotic cells have a plasma membrane ((Figure)), a phospholipid bilayer with embedded proteins that separates the internal contents of the cell from its surrounding environment. A phospholipid is a lipid molecule with two fatty acid chains and a phosphate-containing group. The plasma membrane controls the passage of organic molecules, ions, water, and oxygen into and out of the cell. Wastes (such as carbon dioxide and ammonia) also leave the cell by passing through the plasma membrane.

The eukaryotic plasma membrane is a phospholipid bilayer with proteins and cholesterol embedded in it.


The plasma membrane is composed of a phospholipid bilayer. In the bilayer, the two long hydrophobic tails of phospholipids face toward the center, and the hydrophilic head group faces the exterior. Integral membrane proteins and protein channels span the entire bilayer. Protein channels have a pore in the middle. Peripheral membrane proteins sit on the surface of the phospholipids, and are associated with the phospholipid head groups. On the exterior side of the membrane, carbohydrates are attached to certain proteins and lipids. Filaments of the cytoskeleton line the interior of the membrane.

The plasma membranes of cells that specialize in absorption fold into fingerlike projections that we call microvilli (singular = microvillus); ((Figure)). Such cells typically line the small intestine, the organ that absorbs nutrients from digested food. This is an excellent example of form following function.
People with celiac disease have an immune response to gluten, which is a protein in wheat, barley, and rye. The immune response damages microvilli, and thus, afflicted individuals cannot absorb nutrients. This leads to malnutrition, cramping, and diarrhea. Patients suffering from celiac disease must follow a gluten-free diet.

Microvilli, as they appear on cells lining the small intestine, increase the surface area available for absorption. These microvilli are only on the area of the plasma membrane that faces the cavity from which substances will be absorbed. (credit “micrograph”: modification of work by Louisa Howard)


The left part of this figure is a transmission electron micrograph of microvilli, which appear as long, slender stalks extending from the plasma membrane. The right side illustrates cells containing microvilli. The microvilli cover the surface of the cell facing the interior of the small intestine.

The Cytoplasm

The cytoplasm is the cell’s entire region between the plasma membrane and the nuclear envelope (a structure we will discuss shortly). It is comprised of organelles suspended in the gel-like cytosol, the cytoskeleton, and various chemicals ((Figure)). Even though the cytoplasm consists of 70 to 80 percent water, it has a semi-solid consistency, which comes from the proteins within it. However, proteins are not the only organic molecules in the cytoplasm. Glucose and other simple sugars, polysaccharides, amino acids, nucleic acids, fatty acids, and derivatives of glycerol are also there. Ions of sodium, potassium, calcium, and many other elements also dissolve in the cytoplasm. Many metabolic reactions, including protein synthesis, take place in the cytoplasm.

The Nucleus

Typically, the nucleus is the most prominent organelle in a cell ((Figure)). The nucleus (plural = nuclei) houses the cell’s DNA and directs the synthesis of ribosomes and proteins. Let’s look at it in more detail ((Figure)).

The nucleus stores chromatin (DNA plus proteins) in a gel-like substance called the nucleoplasm. The nucleolus is a condensed chromatin region where ribosome synthesis occurs. We call the nucleus’ boundary the nuclear envelope. It consists of two phospholipid bilayers: an outer and an inner membrane. The nuclear membrane is continuous with the endoplasmic reticulum. Nuclear pores allow substances to enter and exit the nucleus.


The Nuclear Envelope

The nuclear envelope is a double-membrane structure that constitutes the nucleus’ outermost portion ((Figure)). Both the nuclear envelope’s inner and outer membranes are phospholipid bilayers.

The nuclear envelope is punctuated with pores that control the passage of ions, molecules, and RNA between the nucleoplasm and cytoplasm. The nucleoplasm is the semi-solid fluid inside the nucleus, where we find the chromatin and the nucleolus.

Chromatin and Chromosomes

To understand chromatin, it is helpful to first explore chromosomes, structures within the nucleus that are made up of DNA, the hereditary material. You may remember that in prokaryotes, DNA is organized into a single circular chromosome. In eukaryotes, chromosomes are linear structures. Every eukaryotic species has a specific number of chromosomes in the nucleus of each cell. For example, in humans, the chromosome number is 46, while in fruit flies, it is eight.
Chromosomes are only visible and distinguishable from one another when the cell is getting ready to divide. When the cell is in the growth and maintenance phases of its life cycle, proteins attach to chromosomes, and they resemble an unwound, jumbled bunch of threads. We call these unwound protein-chromosome complexes chromatin ((Figure)). Chromatin describes the material that makes up the chromosomes both when condensed and decondensed.

(a) This image shows various levels of chromatin’s organization (DNA and protein). (b) This image shows paired chromosomes. (credit b: modification of work by NIH; scale-bar data from Matt Russell)

Part a: In this illustration, DNA tightly coiled into two thick cylinders is shown in the upper right. A close-up shows how the DNA is coiled around proteins called histones.

Part b: This image shows paired chromosomes.

The Nucleolus

We already know that the nucleus directs the synthesis of ribosomes, but how does it do this? Some chromosomes have sections of DNA that encode ribosomal RNA. A darkly staining area within the nucleus called the nucleolus (plural = nucleoli) aggregates the ribosomal RNA with associated proteins to assemble the ribosomal subunits that are then transported out through the pores in the nuclear envelope to the cytoplasm.

Ribosomes

Ribosomes are the cellular structures responsible for protein synthesis. When we view them through an electron microscope, ribosomes appear either as clusters (polyribosomes) or single, tiny dots that float freely in the cytoplasm. They may be attached to the plasma membrane’s cytoplasmic side or the endoplasmic reticulum’s cytoplasmic side and the nuclear envelope’s outer membrane ((Figure)). Electron microscopy shows us that ribosomes, which are large protein and RNA complexes, consist of two subunits, large and small ((Figure)). Ribosomes receive their “orders” for protein synthesis from the nucleus where the DNA transcribes into messenger RNA (mRNA). The mRNA travels to the ribosomes, which translate the code provided by the sequence of the nitrogenous bases in the mRNA into a specific order of amino acids in a protein. Amino acids are the building blocks of proteins.

A large subunit (top) and a small subunit (bottom) comprise ribosomes. During protein synthesis, ribosomes assemble amino acids into proteins.


The ribosome consists of a small subunit and a large subunit, which is about three times as big as the small one. The large subunit sits on top of the small one. A chain of mRNA threads between the large and small subunits. A protein chain extends from the top of the large subunit.

Because protein synthesis is an essential function of all cells (including enzymes, hormones, antibodies, pigments, structural components, and surface receptors), there are ribosomes in practically every cell. Ribosomes are particularly abundant in cells that synthesize large amounts of protein. For example, the pancreas is responsible for creating several digestive enzymes and the cells that produce these enzymes contain many ribosomes. Thus, we see another example of form following function.

Mitochondria

Scientists often call mitochondria (singular = mitochondrion) the cell’s “powerhouses” or “energy factories” because they are responsible for making adenosine triphosphate (ATP), the cell’s main energy-carrying molecule. ATP represents the cell’s short-term stored energy. Cellular respiration is the process of making ATP using the chemical energy in glucose and other nutrients. In mitochondria, this process uses oxygen and produces carbon dioxide as a waste product. In fact, the carbon dioxide that you exhale with every breath comes from the cellular reactions that produce carbon dioxide as a byproduct.

In keeping with our theme of form following function, it is important to point out that muscle cells have a very high concentration of mitochondria that produce ATP. Your muscle cells need considerable energy to keep your body moving. When your cells don’t get enough oxygen, they do not make much ATP. Instead, producing lactic acid accompanies the small amount of ATP they make in the absence of oxygen.

Mitochondria are oval-shaped, double membrane organelles ((Figure)) that have their own ribosomes and DNA. Each membrane is a phospholipid bilayer embedded with proteins. The inner layer has folds called cristae. We call the area surrounded by the folds the mitochondrial matrix. The cristae and the matrix have different roles in cellular respiration.

This electron micrograph shows a mitochondrion through an electron microscope. This organelle has an outer membrane and an inner membrane. The inner membrane contains folds, called cristae, which increase its surface area. We call the space between the two membranes the intermembrane space, and the space inside the inner membrane the mitochondrial matrix. ATP synthesis takes place on the inner membrane. (credit: modification of work by Matthew Britton; scale-bar data from Matt Russell)


This transmission electron micrograph of a mitochondrion shows an oval outer membrane and an inner membrane with many folds called cristae. Inside the inner membrane is a space called the mitochondrial matrix.

Peroxisomes

Peroxisomes are small, round organelles enclosed by single membranes. They carry out oxidation reactions that break down fatty acids and amino acids. They also detoxify many poisons that may enter the body. (Many of these oxidation reactions release hydrogen peroxide, H2O2, which would be damaging to cells; however, when these reactions are confined to peroxisomes, enzymes safely break down the H2O2 into oxygen and water.) For example, peroxisomes in liver cells detoxify alcohol. Glyoxysomes, which are specialized peroxisomes in plants, are responsible for converting stored fats into sugars. Plant cells contain many different types of peroxisomes that play a role in metabolism, pathogene defense, and stress response, to mention a few.

Vesicles and Vacuoles

Vesicles and vacuoles are membrane-bound sacs that function in storage and transport. Other than the fact that vacuoles are somewhat larger than vesicles, there is a very subtle distinction between them. Vesicle membranes can fuse with either the plasma membrane or other membrane systems within the cell. Additionally, some agents such as enzymes within plant vacuoles break down macromolecules. The vacuole’s membrane does not fuse with the membranes of other cellular components.

Animal Cells versus Plant Cells

At this point, you know that each eukaryotic cell has a plasma membrane, cytoplasm, a nucleus, ribosomes, mitochondria, peroxisomes, and in some, vacuoles, but there are some striking differences between animal and plant cells. While both animal and plant cells have microtubule organizing centers (MTOCs), animal cells also have centrioles associated with the MTOC: a complex we call the centrosome. Animal cells each have a centrosome and lysosomes; whereas, most plant cells do not. Plant cells have a cell wall, chloroplasts and other specialized plastids, and a large central vacuole; whereas, animal cells do not.

The Centrosome

The centrosome is a microtubule-organizing center found near the nuclei of animal cells. It contains a pair of centrioles, two structures that lie perpendicular to each other ((Figure)). Each centriole is a cylinder of nine triplets of microtubules.

The centrosome consists of two centrioles that lie at right angles to each other. Each centriole is a cylinder comprised of nine triplets of microtubules. Nontubulin proteins (indicated by the green lines) hold the microtubule triplets together.


The centrosome (the organelle where all microtubules originate) replicates itself before a cell divides, and the centrioles appear to have some role in pulling the duplicated chromosomes to opposite ends of the dividing cell. However, the centriole’s exact function in cell division isn’t clear, because cells that have had the centrosome removed can still divide, and plant cells, which lack centrosomes, are capable of cell division.

Lysosomes

Animal cells have another set of organelles that most plant cells do not: lysosomes. The lysosomes are the cell’s “garbage disposal.” In plant cells, the digestive processes take place in vacuoles. Enzymes within the lysosomes aid in breaking down proteins, polysaccharides, lipids, nucleic acids, and even worn-out organelles. These enzymes are active at a much lower pH than the cytoplasm’s. Therefore, the pH within lysosomes is more acidic than the cytoplasm’s pH. Many reactions that take place in the cytoplasm could not occur at a low pH, so again, the advantage of compartmentalizing the eukaryotic cell into organelles is apparent.

The Cell Wall

If you examine (Figure), the plant cell diagram, you will see a structure external to the plasma membrane. This is the cell wall, a rigid covering that protects the cell, provides structural support, and gives shape to the cell. Fungal and some protistan cells also have cell walls. While the prokaryotic cell walls’ chief component is peptidoglycan, the major organic molecule in the plant (and some protists’) cell wall is cellulose ((Figure)), a polysaccharide comprised of glucose units. Have you ever noticed that when you bite into a raw vegetable, like celery, it crunches? That’s because you are tearing the celery cells’ rigid cell walls with your teeth.

Cellulose is a long chain of β-glucose molecules connected by a 1-4 linkage. The dashed lines at each end of the figure indicate a series of many more glucose units. The size of the page makes it impossible to portray an entire cellulose molecule.


This illustration shows three glucose subunits that are attached together. Dashed lines at each end indicate that many more subunits make up an entire cellulose fiber. Each glucose subunit is a closed ring composed of carbon, hydrogen, and oxygen atoms.

Chloroplasts

Like the mitochondria, chloroplasts have their own DNA and ribosomes, but chloroplasts have an entirely different function. Chloroplasts are plant cell organelles that carry out photosynthesis. Photosynthesis is the series of reactions that use carbon dioxide, water, and light energy to make glucose and oxygen. This is a major difference between plants and animals. Plants (autotrophs) are able to make their own food, like sugars, while animals (heterotrophs) must ingest their food.

Like mitochondria, chloroplasts have outer and inner membranes, but within the space enclosed by a chloroplast’s inner membrane is a set of interconnected and stacked fluid-filled membrane sacs we call thylakoids ((Figure)). Each thylakoid stack is a granum (plural = grana). We call the fluid enclosed by the inner membrane that surrounds the grana the stroma.

The chloroplast has an outer membrane, an inner membrane, and membrane structures – thylakoids that are stacked into grana. We call the space inside the thylakoid membranes the thylakoid space. The light harvesting reactions take place in the thylakoid membranes, and sugar synthesis takes place in the fluid inside the inner membrane, which we call the stroma. Chloroplasts also have their own genome, which is contained on a single circular chromosome.


This illustration shows a chloroplast, which has an outer membrane and an inner membrane. The space between the outer and inner membranes is called the intermembrane space. Inside the inner membrane are flat, pancake-like structures called thylakoids. The thylakoids form stacks called grana. The liquid inside the inner membrane is called the stroma, and the space inside the thylakoids is called the thylakoid space.

The chloroplasts contain a green pigment, chlorophyll, which captures the light energy that drives the reactions of photosynthesis. Like plant cells, photosynthetic protists also have chloroplasts. Some bacteria perform photosynthesis, but their chlorophyll is not relegated to an organelle.

Evolution Connection

Endosymbiosis

We have mentioned that both mitochondria and chloroplasts contain DNA and ribosomes. Have you wondered why? Strong evidence points to endosymbiosis as the explanation.

Symbiosis is a relationship in which organisms from two separate species depend on each other for their survival. Endosymbiosis (endo- = “within”) is a mutually beneficial relationship in which one organism lives inside the other. Endosymbiotic relationships abound in nature. We have already mentioned that microbes that produce vitamin K live inside the human gut. This relationship is beneficial for us because we are unable to synthesize vitamin K. It is also beneficial for the microbes because they are protected from other organisms and from drying out, and they receive abundant food from the environment of the large intestine.

Scientists have long noticed that bacteria, mitochondria, and chloroplasts are similar in size. We also know that bacteria have DNA and ribosomes, just like mitochondria and chloroplasts. Scientists believe that host cells and bacteria formed an endosymbiotic relationship when the host cells ingested both aerobic and autotrophic bacteria (cyanobacteria) but did not destroy them. Through many millions of years of evolution, these ingested bacteria became more specialized in their functions, with the aerobic bacteria becoming mitochondria and the autotrophic bacteria becoming chloroplasts.

The Central Vacuole

Previously, we mentioned vacuoles as essential components of plant cells. If you look at (Figure)b, you will see that plant cells each have a large central vacuole that occupies most of the cell’s area. The central vacuole plays a key role in regulating the cell’s concentration of water in changing environmental conditions. Have you ever noticed that if you forget to water a plant for a few days, it wilts? That’s because as the water concentration in the soil becomes lower than the water concentration in the plant, water moves out of the central vacuoles and cytoplasm. As the central vacuole shrinks, it leaves the cell wall unsupported. This loss of support to the plant’s cell walls results in the wilted appearance.

The central vacuole also supports the cell’s expansion. When the central vacuole holds more water, the cell becomes larger without having to invest considerable energy in synthesizing new cytoplasm.

Section Summary

Like a prokaryotic cell, a eukaryotic cell has a plasma membrane, cytoplasm, and ribosomes, but a eukaryotic cell is typically larger than a prokaryotic cell, has a true nucleus (meaning a membrane surrounds its DNA), and has other membrane-bound organelles that allow for compartmentalizing functions. The plasma membrane is a phospholipid bilayer embedded with proteins. The nucleus’s nucleolus is the site of ribosome assembly. We find ribosomes either in the cytoplasm or attached to the cytoplasmic side of the plasma membrane or endoplasmic reticulum. They perform protein synthesis. Mitochondria participate in cellular respiration. They are responsible for the majority of ATP produced in the cell. Peroxisomes hydrolyze fatty acids, amino acids, and some toxins. Vesicles and vacuoles are storage and transport compartments. In plant cells, vacuoles also help break down macromolecules.

Animal cells also have a centrosome and lysosomes. The centrosome has two bodies perpendicular to each other, the centrioles, and has an unknown purpose in cell division. Lysosomes are the digestive organelles of animal cells.

Plant cells and plant-like cells each have a cell wall, chloroplasts, and a central vacuole. The plant cell wall, whose primary component is cellulose, protects the cell, provides structural support, and gives the cell shape. Photosynthesis takes place in chloroplasts. The central vacuole can expand without having to produce more cytoplasm.

Art Connections

(Figure) If the nucleolus were not able to carry out its function, what other cellular organelles would be affected?

Review Questions

Which of the following is surrounded by two phospholipid bilayers?

  1. the ribosomes
  2. the vesicles
  3. the cytoplasm
  4. the nucleoplasm

Peroxisomes got their name because hydrogen peroxide is:

  1. used in their detoxification reactions
  2. produced during their oxidation reactions
  3. incorporated into their membranes
  4. a cofactor for the organelles’ enzymes

In plant cells, the function of the lysosomes is carried out by __________.

  1. vacuoles
  2. peroxisomes
  3. ribosomes
  4. nuclei

Which of the following is both in eukaryotic and prokaryotic cells?

  1. nucleus
  2. mitochondrion
  3. vacuole
  4. ribosomes

Tay-Sachs disease is a genetic disorder that results in the destruction of neurons due to a buildup of sphingolipids in the cells. Which organelle is malfunctioning in Tay-Sachs?

  1. lysosome
  2. endoplasmic reticulum
  3. peroxisome
  4. mitochondria

Free Response

You already know that ribosomes are abundant in red blood cells. In what other cells of the body would you find them in great abundance? Why?

What are the structural and functional similarities and differences between mitochondria and chloroplasts?

Why are plasma membranes arranged as a bilayer rather than a monolayer?

Glossary

cell wall
rigid cell covering comprised of various molecules that protects the cell, provides structural support, and gives shape to the cell
central vacuole
large plant cell organelle that regulates the cell’s storage compartment, holds water, and plays a significant role in cell growth as the site of macromolecule degradation
centrosome
region in animal cells made of two centrioles that serves as an organizing center for microtubules
chlorophyll
green pigment that captures the light energy that drives the light reactions of photosynthesis
chloroplast
plant cell organelle that carries out photosynthesis
chromatin
protein-DNA complex that serves as the chromosomes’ building material
chromosome
structure within the nucleus that comprises chromatin that contains DNA, the hereditary material
cytoplasm
entire region between the plasma membrane and the nuclear envelope, consisting of organelles suspended in the gel-like cytosol, the cytoskeleton, and various chemicals
cytosol
the cytoplasm’s gel-like material in which cell structures are suspended
eukaryotic cell
cell that has a membrane-bound nucleus and several other membrane-bound compartments or sacs
lysosome
organelle in an animal cell that functions as the cell’s digestive component; it breaks down proteins, polysaccharides, lipids, nucleic acids, and even worn-out organelles
mitochondria
(singular = mitochondrion) cellular organelles responsible for carrying out cellular respiration, resulting in producing ATP, the cell’s main energy-carrying molecule
nuclear envelope
double-membrane structure that constitutes the nucleus’ outermost portion
nucleolus
darkly staining body within the nucleus that is responsible for assembling ribosome subunits
nucleoplasm
semi-solid fluid inside the nucleus that contains the chromatin and nucleolus
nucleus
cell organelle that houses the cell’s DNA and directs ribosome and protein synthesis
organelle
compartment or sac within a cell
peroxisome
small, round organelle that contains hydrogen peroxide, oxidizes fatty acids and amino acids, and detoxifies many poisons
plasma membrane
phospholipid bilayer with embedded (integral) or attached (peripheral) proteins, and separates the cell’s internal content from its surrounding environment
ribosome
cellular structure that carries out protein synthesis
vacuole
membrane-bound sac, somewhat larger than a vesicle, which functions in cellular storage and transport
vesicle
small, membrane-bound sac that functions in cellular storage and transport; its membrane is capable of fusing with the plasma membrane and the membranes of the endoplasmic reticulum and Golgi apparatus