Mendoza, Alyssa Marie G.
Properties of Life
All living organisms share several key characteristics or functions: order, sensitivity or response
to the environment, reproduction, growth and development, regulation, homeostasis, and energy
processing. When viewed together, these characteristics serve to define life.
Order
Organisms are highly organized, coordinated structures that consist of one or more cells. Even
very simple, single-celled organisms are remarkably complex: inside each cell, atoms make up
molecules; these in turn make up cell organelles and other cellular inclusions.
In multicellular organisms (Figure 1), similar cells form tissues. Tissues, in turn, collaborate to
create organs (body structures with a distinct function). Organs work together to form organ
systems.
Sensitivity or Response to Stimuli
Organisms respond to diverse stimuli. For example, plants can bend toward a source of light,
climb on fences and walls, or respond to touch (Figure 2).
Even tiny bacteria can move toward or away from chemicals (a process called chemotaxis) or
light (phototaxis). Movement toward a stimulus is considered a positive response, while
movement away from a stimulus is considered a negative response.
Reproduction
Single-celled organisms reproduce by first duplicating their DNA, and then dividing it equally as
the cell prepares to divide to form two new cells. Multicellular organisms often produce
specialized reproductive germline cells that will form new individuals. When reproduction occurs,
genes containing DNA are passed along to an organism’s offspring. These genes ensure that the
offspring will belong to the same species and will have similar characteristics, such as size and
shape.
Growth and Development
Organisms grow and develop following specific instructions coded for by their genes. These
genes provide instructions that will direct cellular growth and development, ensuring that a
species’ young (Figure 3) will grow up to exhibit many of the same characteristics as its parents.
Regulation
Even the smallest organisms are complex and require multiple regulatory mechanisms to
coordinate internal functions, respond to stimuli, and cope with environmental stresses. Two
examples of internal functions regulated in an organism are nutrient transport and blood flow.
Organs (groups of tissues working together) perform specific functions, such as carrying oxygen
throughout the body, removing wastes, delivering nutrients to every cell, and cooling the body.
Homeostasis
In order to function properly, cells need to have appropriate conditions such as proper
temperature, pH, and appropriate concentration of diverse chemicals. These conditions may,
however, change from one moment to the next. Organisms are able to maintain internal
conditions within a narrow range almost constantly, despite environmental changes,
through homeostasis (literally, “steady state”)—the ability of an organism to maintain constant
internal conditions. For example, an organism needs to regulate body temperature through a
process known as thermoregulation. Organisms that live in cold climates, such as the polar bear
(Figure 4), have body structures that help them withstand low temperatures and conserve body
heat. Structures that aid in this type of insulation include fur, feathers, blubber, and fat. In hot
climates, organisms have methods (such as perspiration in humans or panting in dogs) that help
them to shed excess body heat.
Energy Processing
All organisms use a source of energy for their metabolic activities. Some organisms capture
energy from the sun and convert it into chemical energy in food (photosynthesis); others use
chemical energy in molecules they take in as food (cellular respiration).
Levels of Organization of Living Things
Living things are highly organized and structured, following a hierarchy that can be examined on
a scale from small to large. The atomis the smallest and most fundamental unit of matter. It
consists of a nucleus surrounded by electrons. Atoms form molecules. A molecule is a chemical
structure consisting of at least two atoms held together by one or more chemical bonds. Many
molecules that are biologically important are macromolecules, large molecules that are typically
formed by polymerization (a polymer is a large molecule that is made by combining smaller units
called monomers, which are simpler than macromolecules). An example of a macromolecule is
deoxyribonucleic acid (DNA) (Figure 6), which contains the instructions for the structure and
functioning of all living organisms.
Some cells contain aggregates of macromolecules surrounded by membranes; these are
called organelles. Organelles are small structures that exist within cells. Examples of organelles
include mitochondria and chloroplasts, which carry out indispensable functions: mitochondria
produce energy to power the cell, while chloroplasts enable green plants to utilize the energy in
sunlight to make sugars. All living things are made of cells; the cell itself is the smallest
fundamental unit of structure and function in living organisms. (This requirement is why viruses
are not considered living: they are not made of cells. To make new viruses, they have to invade
and hijack the reproductive mechanism of a living cell; only then can they obtain the materials
they need to reproduce.) Some organisms consist of a single cell and others are multicellular.
Cells are classified as prokaryotic or eukaryotic. Prokaryotes are single-celled or colonial
organisms that do not have membrane-bound nuclei or organelles; in contrast, the cells
of eukaryotes do have membrane-bound organelles and a membrane-bound nucleus.
In larger organisms, cells combine to make tissues, which are groups of similar cells carrying out
similar or related functions. Organs are collections of tissues grouped together performing a
common function. Organs are present not only in animals but also in plants. An organ system is
a higher level of organization that consists of functionally related organs. Mammals have many
organ systems. For instance, the circulatory system transports blood through the body and to and
from the lungs; it includes organs such as the heart and blood vessels. Organisms are individual
living entities. For example, each tree in a forest is an organism. Single-celled prokaryotes and
single-celled eukaryotes are also considered organisms and are typically referred to as
microorganisms.
All the individuals of a species living within a specific area are collectively called a population.
For example, a forest may include many pine trees. All of these pine trees represent the
population of pine trees in this forest. Different populations may live in the same specific area. For
example, the forest with the pine trees includes populations of flowering plants and also insects
and microbial populations. A community is the sum of populations inhabiting a particular area.
For instance, all of the trees, flowers, insects, and other populations in a forest form the forest’s
community. The forest itself is an ecosystem. An ecosystem consists of all the living things in a
particular area together with the abiotic, non-living parts of that environment such as nitrogen in
the soil or rain water. At the highest level of organization (Figure 7), the biosphere is the
collection of all ecosystems, and it represents the zones of life on earth. It includes land, water,
and even the atmosphere to a certain extent.
Cell Theory is one of the basic principles of biology. Credit for the formulation of this theory is
given to German scientists Theodor Schwann, Matthias Schleiden, and Rudolph Virchow.
The Cell Theory states:
 All living organisms are composed of cells. They may be unicellular or
multicellular.
 The cell is the basic unit of life.
 Cells arise from pre-existing cells. (They are not derived from spontaneous
generation.)
The modern version of the Cell Theory includes the ideas that:
 Energy flow occurs within cells.
 Heredity information (DNA) is passed on from cell to cell.
 All cells have the same basic chemical composition.
In addition to the cell theory, the gene theory, evolution, homeostasis, and the laws of
thermodynamics form the basic principles that are the foundation for the study of life.
What Are Cells?
Cells are the simplest unit of matter that is living. The two primary kinds of cells
are eukaryotic cells, which have a true nucleuscontaining DNA and prokaryotic cells, which
have no true nucleus. In prokaryotic cells, the DNA is coiled up in a region called the nucleoid.
Cell Basics
All living organisms in the kingdoms of life are composed of and depend on cells to function
normally. Not all cells, however, are alike. There are two primary types of cells: eukaryotic and
prokaryotic cells. Examples of eukaryotic cells include animal cells, plant cells, and fungal
cells. Prokaryotic cells include bacteria and archaeans.
Cells contain organelles, or tiny cellular structures, that carry out specific functions necessary for
normal cellular operation. Cells also contain DNA (deoxyribonucleic acid) and RNA (ribonucleic
acid), the genetic information necessary for directing cellular activities.
Cell Reproduction
Eukaryotic cells grow and reproduce through a complex sequence of events called the cell cycle.
At the end of the cycle, cells will divide either through the processes of mitosis or meiosis.
Somatic cells replicate through mitosis and sex cells reproduce via meiosis. Prokaryotic cells
reproduce commonly through a type of asexual reproduction called binary fission. Higher
organisms are also capable of asexual reproduction. Plants, algae, and fungi reproduce through
the formation of reproductive cells called spores. Animal organisms can reproduce asexually
through processes such as budding, fragmentation, regeneration, and parthenogenesis.
Cell Processes - Cellular Respiration and Photosynthesis
Cells perform a number of important processes that are necessary for the survival of an
organism. Cells undergo the complex process of cellular respiration in order to obtain energy
stored in the nutrients consumed. Photosynthetic organisms including plants, algae,
and cyanobacteria are capable of photosynthesis. In photosynthesis, light energy from the sun is
converted to glucose. Glucose is the energy source used by photosynthetic organisms and other
organisms that consume photosynthetic organisms.
Cell Processes - Endocytosis and Exocytosis
Cells also perform the active transport processes of endocytosis and exocytosis. Endocytosis is
the process of internalizing and digesting substances, such as seen
with macrophages and bacteria. The digested substances are expelled through exocytosis.
These processes also allow for molecule transportation between cells.
Cell Processes - Cell Migration
Cell migration is a process that is vital for the development of tissues and organs. Cell movement
is also required for mitosis and cytokinesis to occur. Cell migration is made possible by
interactions between motor enzymes and cytoskeleton microtubules.
Cell Processes - DNA Replication and Protein Synthesis
The cell process of DNA replication is an important function that is needed for several
processes including chromosome synthesis and cell division to occur. DNA
transcription and RNA translation make the process of protein synthesis possible.
2.3 Cell structure and function (ESG4S)
Cell theory (ESG4T)
The cell theory developed in 1839 by microbiologists Schleiden and Schwann describes the
properties of cells. It is an explanation of the relationship between cells and living things. The
theory states that:
 all living things are made of cells and their products.
 new cells are created by old cells dividing into two.
 cells are the basic building blocks of life.
The cell theory applies to all living things, however big or small. The modern understanding of cell
theory extends the concepts of the original cell theory to include the following:
 The activity of an organism depends on the total activity of independent cells.
Mendoza, Alyssa Marie G.
 
Energy flow occurs in cells through the breakdown of carbohydrates by
respiration.
 Cells contain the information necessary for the creation of new cells. This
information is known as 'hereditary information' and is contained within DNA.
 The contents of cells from similar species are basically the same.
DNA (the hereditary information of cells) is passed from 'parent' cells to 'daughter' cells during cell
division. You will learn more about this in the following chapter: Cell division.
Cells are the smallest form of life; the functional and structural units of all living things. Your body
contains several billion cells, organised into over 200 major types, with hundreds of cell-specific
functions.
Some functions performed by cells are so vital to the existence of life that all cells perform them
(e.g. cellular respiration). Others are highly specialised (e.g. photosynthesis).
Figure 2.9 shows a two-dimensional drawing of an animal cell. The diagram shows the structures
visible within a cell at high magnification. The structures form the ultrastructure of the cell.
Figure 2.9: Diagram of the cell ultrastructure of an
animal cell.
We will now look at some of the basic cell structures
and organelles in animal and plant cells.
Cell wall (ESG4V)
The cell wall is a rigid non-living layer that is found outside the cell membrane and surrounds the
cell. Plants, bacteria and fungi all have cell walls. In plants, the wall is comprised of cellulose. It
consists of three layers that help support the plant. These layers include the middle lamella, the
primary cell wall and the secondary cell wall.
Middle lamella: Separates one cell from another. It is a thin membranous layer on the outside of
the cell and is made of a sticky substance called pectin.
Primary cell wall: Is on the inside of the middle lamella and is mainly composed of cellulose.
Secondary cell wall: Lies alongside the cell membrane. It is is made up of a thick and tough
layer of cellulose which is held together by a hard, waterproof substance called lignin. It is only
found in cells which provide mechanical support in plants.
The human body cannot break down the cellulose in
cell walls because we do not produce the enzyme
cellulase.
Figure 2.10: Scanning electron microscope
micrographs of diatoms showing the external
appearances of the cell wall. Scale bar: A, B, D: 10
um, C 20 um
Functions of the cell wall

The main function of the wall is to protect the inner parts of the plant cell, it gives
plant cells a more uniform and regular shape and provides support for the plant body.
 The cell wall is completely permeable to water and mineral salts which allows
distribution of nutrients throughout the plant.
Image of cyanobacterial cells forming a filament composed of dissimilar cell types. Most of the
The openings in the cell wall are called plasmodesmata which contain strands of cells are small, but one is round and different in morphology. This larger cell is a nitrogen-fixing
cytoplasm that connect adjacent cells. This allows cells to interact with one another, allowing cell.
molecules to travel between plant cells.
Cell membrane (ESG4W)
Components of prokaryotic cells
The cell membrane, also called the plasma membrane, physically separates the intracellular
space (inside the cell) from the extracellular environment (outside the cell). All plant and animal
cells have cell membranes. The cell membrane surrounds and protects the cytoplasm.
There are some key ingredients that a cell needs in order to be a cell, regardless of whether it is
Cytoplasm is part of the protoplasm and is the living component of the cell.
prokaryotic or eukaryotic. All cells share four key components:
1. The plasma membrane is an outer covering that separates the cell’s interior from its surrounding
The cell membrane is composed of a double layer (bilayer) of special lipids (fats) environment.
called phospholipids. Phospholipids consist of a hydrophilic (water-loving) head and 2. Cytoplasm consists of the jelly-like cytosol inside the cell, plus the cellular structures suspended
a hydrophobic (water-fearing) tail. The hydrophobic head of the phospholipid is polar (charged) in it. In eukaryotes, cytoplasm specifically means the region outside the nucleus but inside the
and can therefore dissolve in water. The hydrophobic tail is non-polar (uncharged), and cannot plasma membrane.
dissolve in water. 3. DNA is the genetic material of the cell.
4. Ribosomes are molecular machines that synthesize proteins.
Despite these similarities, prokaryotes and eukaryotes differ in a number of important ways.
A prokaryote is a simple, single-celled organism that lacks a nucleus and membrane-bound
The lipid bilayer forms spontaneously due to the properties of the phospholipid molecules. In an organelles. We’ll talk more about the nucleus and organelles in the next article on eukaryotic
aqueous environment, the polar heads try to form hydrogen bonds with the water, while the non- cells, but the main thing to keep in mind for now is that prokaryotic cells are not divided up on the
polar tails try to escape from the water. The problem is solved by the formation of a bilayer inside by membrane walls, but consist instead of a single open space.
because the hydrophilic heads can point outwards and from hydrogen bonds with water, and the
hydrophobic tails point towards one another and are 'protected' from the water molecules (Figure
2.11. The majority of prokaryotic \text{DNA}DNAD, N, A is found in a central region of the cell called
the nucleoid, and it typically consists of a single large loop called a circular chromosome. The
nucleoid and some other frequently seen features of prokaryotes are shown in the diagram below
of a cut-away of a rod-shaped bacterium.
Figure 2.11: The lipid bilayer showing the arrangement of phospholipids, containing hydrophilic,
polar heads and hydrophobic,
Image of a typical prokaryotic cell, with different portions
non-polar tails.
of the cell labeled.
_Image credit: modified from "Prokaryotic cells: Figure
Recall the structure of lipid 1" by OpenStax College, Biology, CC BY 3.0_
molecules from the previous
chapter on the chemistry of
life. Bacteria are very diverse in form, so not every type of
All the exchanges between bacterium will have all of the features shown in the
the cell and its environment diagram.
have to pass through the cell
membrane. The cell
membrane is selectively permeable to ions (e.g. hydrogen, sodium), small molecules (oxygen, Most bacteria are, however, surrounded by a rigid cell wall
made out of peptidoglycan, a polymer composed of linked carbohydrates
carbon dioxide) and larger molecules (glucose and amino acids) and controls the movement of
and small proteins. The cell wall provides an extra layer of protection, helps the cell maintain its
substances in and out of the cells. The cell membrane performs many important functions within
shape, and prevents dehydration. Many bacteria also have an outermost layer of carbohydrates
the cell such as osmosis, diffusion, transport of nutrients into the cell, processes of ingestion and
called the capsule. The capsule is sticky and helps the cell attach to surfaces in its environment.
secretion. The cell membrane is strong enough to provide the cell with mechanical support and
flexible enough to allow cells to grow and move.
Some bacteria also have specialized structures found on the cell surface, which may help them
move, stick to surfaces, or even exchange genetic material with other bacteria. For
instance, flagella are whip-like structures that act as rotary motors to help bacteria move.
Fimbriae are numerous, hair-like structures that are used for attachment to host cells and other
surfaces. Bacteria may also have rod-like structures known as pili, which come in different
Introduction
varieties. For instance, some types of pili allow a bacterium to transfer \text{DNA}DNAD, N,
A molecules to other bacteria, while others are involved in bacterial locomotion—helping the
bacterium move.
Take a moment and look at yourself. How many organisms do you see? Your first thought might
be that there's just one: yourself. However, if you were to look closer, at the surface of your skin
or inside your digestive tract, you would see that there are actually many organisms living there. [Are fimbriae considered pili?]
That’s right - you are home to around 100 trillion bacterial cells!
^1start superscript, 1, end superscript
This means that your body is actually an ecosystem. It also means that you—for some definition
of the word you—actually consist of both of the major types of cells: prokaryotic and eukaryotic.
Archaea may
also have most
All cells fall into one of these two broad categories. Only the single-celled organisms of the of these cell
domains Bacteria and Archaea are classified as prokaryotes—pro means before and kary means surface
nucleus. Animals, plants, fungi, and protists are all eukaryotes—eu means true—and are made features, but
up of eukaryotic cells. Often, though—as in the case of we humans—there are some prokaryotic their versions of
friends hanging around. a particular
feature are
typically
[Are all prokaryotes single-celled?] different from
those of
\text N_2N, start subscript, 2, end subscript bacteria. For
instance,
although
archaea also
have a cell wall,
it's not made
out of
Mendoza, Alyssa Marie G.
peptidoglycan—although it does contain carbohydrates and proteins.
Cell size
Typical prokaryotic cells range from 0.1 to 5.0 micrometers (μm) in diameter and are significantly
smaller than eukaryotic cells, which usually have diameters ranging from 10 to 100 μm.
The figure below shows the sizes of prokaryotic, bacterial, and eukaryotic, plant and animal, cells
as well as other molecules and organisms on a logarithmic scale. Each unit of increase in a
logarithmic scale represents a 10-fold increase in the quantity being measured, so these are big
size differences we’re talking about!
Graph showing the relative sizes of items from, in order, atoms to proteins to viruses to bacteria to
animal cells to chicken eggs to humans.
_Image credit: "Prokaryotic cells: FIgure 2" by OpenStax College, Biology, CC BY 3.0_
With a few cool exceptions—check out the single-celled seaweed Caulerpa—cells must remain
fairly small, regardless of whether they’re prokaryotic or eukaryotic. Why should this be the case?
The basic answer is that as cells become larger, it gets harder for them to exchange enough
nutrients and wastes with their environment. To see how this works, let’s look at a cell’s surface-
area-to-volume ratio.
Suppose, for the sake of keeping things simple, that we have a cell that’s shaped like a cube.
Some plant cells are, in fact, cube-shaped. If the length of one of the cube’s sides is lll, the
surface area of the cube will be 6l^26l26, l, start superscript, 2, end superscript, and the volume
of the cube will be l^3l3l, start superscript, 3, end superscript. This means that as lll gets bigger,
the surface area will increase quickly since it changes with the square of lll. The volume,
however, will increase even faster since it changes with the cube of lll.
Thus, as a cell gets bigger, its surface-area-to-volume ratio drops. For example, the cube-shaped
cell on the left has a volume of 1 mm^33start superscript, 3, end superscript and a surface area
of 6 mm^22start superscript, 2, end superscript with a surface-area-to-volume ratio of six to one,
whereas the cube-shaped cell on the right has a volume of 8 mm^33start superscript, 3, end
superscript and a surface area of 24 mm^22start superscript, 2, end superscriptwith a surface
area-to-volume ratio of three to one.
Image of two cubes of different sizes. The cube on
the left has 1 mm sides, while the cube on the
right has 2 mm sides.
_Image credit: modified from "Prokaryotic
cells: FIgure 3" by OpenStax College,
Biology, CC BY 3.0_
Surface-area-to-volume ratio is important because the plasma membrane is the cell’s interface
with the environment. If the cell needs to take up nutrients, it must do so across the membrane,
and if it needs to eliminate wastes, the membrane is again its only route.
Each patch of membrane can exchange only so much of a given substance in a given period of
time – for instance, because it contains a limited number of channels. If the cell grows too large,
its membrane will not have enough exchange capacity (surface area, square function) to support
the rate of exchange required for its increased metabolic activity (volume, cube function).
The surface-area-to-volume problem is just one of a related set of difficulties posed by large cell
size. As cells get larger, it also takes longer to transport materials inside of them. These
considerations place a general upper limit on cell size, with eukaryotic cells being able to exceed
prokaryotic cells thanks to their structural and metabolic features—which we’ll explore in the next
section.
Some cells also use geometric tricks to get around the surface-area-to-volume problem.
For instance, some cells are long and thin or have many protrusions from their surface,
features that increase surface area relative to volume^22start superscript, 2, end
superscript.
Introduction
What would it be like to live in a one-room cabin? Well, things would probably be pretty simple.
You would eat, sleep, work, and relax in a single room—which might be a bit cramped, but would
certainly make cleaning the house a snap!
Prokaryotic cells, the simple cells of organisms like bacteria, are sometimes compared to one-
room cabins: they don't have internal membranes, so they’re like a single room with no walls to
Diagram of an animal cell with components lettered.
carve it up^11start superscript, 1, end superscript. If we extend this analogy to eukaryotic cells,
the more complex cells that make up plants, fungi, and animals, we'll find that they're a definite
step upward in the real estate market. Image modified from OpenStax Biology.
Just as a large family home is split into many rooms with different purposes (bedrooms, Diagram of a typical plant cell:
bathrooms, kitchen, living room, etc.), so eukaryotic cells contain a variety of different
compartments with specialized functions, neatly separated from one another by layers of
membrane. This organization lets each compartment maintain its own conditions, the ones it
needs to carry out its job.
For instance, compartments called lysosomes, which act as recycling centers for the cell, must
maintain an acidic pH in order to dispose of cellular waste. Similarly, structures
called peroxisomes carry out chemical reactions called oxidation reactions and produce
hydrogen peroxide, both of which would damage the cell if they weren’t safely stored away in
their own “room.”
The ability to maintain different environments inside a single cell allows eukaryotic cells to carry
out complex metabolic reactions that prokaryotes cannot. In fact, it’s a big part of the reason why
eukaryotic cells can grow to be many times larger than prokaryotic ones.
Prokaryotic vs. eukaryotic cells
Diagram of a plant cell with components labeled.
What are the key features of eukaryotic cells? Unlike prokaryotic cells, eukaryotic cells have:
Image modified from OpenStax Biology.
1. A membrane-bound nucleus, a central cavity surrounded by membrane that houses the cell’s
genetic material.
Introduction
2. A number of membrane-bound organelles, compartments with specialized functions that float in
the cytosol. (Organelle means “little organ,” and this name reflects that the organelles, like the
organs of our body, have unique functions as part of a larger system.)
3. Multiple linear chromosomes, as opposed to the single circular chromosome of a prokaryote. What’s a cell? Well, on some level, it's a bag of goo. The plasma membrane—the outer boundary
Eukaryotic cells are much more complicated than those of prokaryotes. They are packed with a of the cell—is the bag, and the cytoplasm is the goo.
fascinating array of subcellular structures that play important roles in energy balance,
metabolism, and gene expression.
Of course, a cell is ever so much more than just a bag of goo. It's a complex, highly organized
unit, the basic building block of all living things. And the plasma membrane and cytoplasm are
In the articles and videos that follow, we’ll take a tour through eukaryotic plant and animal cells, actually pretty sophisticated.
exploring the unique structures they contain and the role that each structure plays in the life of the
cell.
The membrane is a delicate, two-layered structure of lipids and proteins, and it controls what can
enter and exit the cell. Similarly, the cytoplasm of a eukaryotic cell consists not only of cytosol—a
Already know what part of the cell you want to visit? Use the list below to jump to your region of gel-like substance made up of water, ions, and macromolecules—but also of organelles and the
interest: structural proteins that make up the cytoskeleton, or "skeleton of the cell."
In this article, we’ll take a closer look at the plasma membrane and cytoplasm.
 Plasma membrane and cytoplasm
 Nucleus and ribosomes
 Endomembrane system
 Mitochondria, chloroplasts, and The plasma membrane
peroxisomes
 Cytoskeleton
Both prokaryotic and eukaryotic cells have a plasma membrane, a double layer of lipids that
 Extracellular matrix and cell wall separates the cell interior from the outside environment. This double layer consists largely of
specialized lipids called phospholipids.
 Cell junctions
A phospholipid is made up of a hydrophilic, water-loving, phosphate head, along with two
hydrophobic, water-fearing, fatty acid tails. Phospholipids spontaneously arrange themselves in a
Diagram of a typical animal cell: double-layered structure with their hydrophobic tails pointing inward and their hydrophilic heads
facing outward. This energetically favorable two-layer structure, called a phospholipid bilayer, is
found in many biological membranes.
Mendoza, Alyssa Marie G.
[Close-up of a phospholipid]
Phospholipid, showing hydrophilic
head with phosphate group and
hydrophobic fatty acid tails
As shown below, proteins are also an important
component of the plasma membrane. Some of them
pass all the way through the membrane, serving as
channels or signal receptors, while others are just
attached at the edge. Different types of lipids, such as
cholesterol, may also be found in the cell membrane and
affect its fluidity.
An image of plasma membrane shows the phospholipid bilayer, embedded proteins, and
cholesterol molecules. The membrane separates the extracellular space, outside of the cell, from
the cytosol inside the cell.
Image
credit: modified from OpenStax Biology
The plasma membrane is the border between the interior and exterior of a cell. As such, it
controls passage of various molecules—including sugars, amino acids, ions, and water—into and
out of the cell. How easily these molecules can cross the membrane depends on their size and
polarity. Some small, nonpolar molecules, such as oxygen, can pass directly through the
phospholipid portion of the membrane. Larger and more polar, hydrophilic, molecules, such as
amino acids, must instead cross the membrane by way of protein channels, a process that is
often regulated by the cell. You can learn more about cellular transport in the membranes and
transport section.
The surface area of the plasma membrane limits the exchange of materials between a cell and its
environment. Some cells are specialized in the exchange of wastes or nutrients and have
modifications to increase the area of the plasma membrane. For instance, the membranes of
some nutrient-absorbing cells are folded into fingerlike projections called microvilli,
singular, microvillus. Cells with microvilli cover the inside surface of the small intestine, the
organ that absorbs nutrients from digested food. The microvilli help intestinal cells maximize their
absorption of nutrients from food by increasing plasma membrane surface area.
[Microvilli and celiac disease]
Diagram and micrograph of intestinal cells, showing the protruding "fingers" of plasma
membrane—called microvilli—that contact the fluid inside the small intestine.
Image credit: OpenStax Biology. Micrograph is a modification of work by Louisa Howard.
The cytoplasm
The part of the cell referred to as cytoplasm is slightly different in eukaryotes and prokaryotes. In
eukaryotic cells, which have a nucleus, the cytoplasm is everything between the plasma
membrane and the nuclear envelope. In prokaryotes, which lack a nucleus, cytoplasm simply
means everything found inside the plasma membrane.
One major component of the cytoplasm in both prokaryotes and eukaryotes is the gel-
like cytosol, a water-based solution that contains ions, small molecules, and macromolecules. In
eukaryotes, the cytoplasm also includes membrane-bound organelles, which are suspended in
the cytosol. The cytoskeleton, a network of fibers that supports the cell and gives it shape, is also
part of the cytoplasm and helps to organize cellular components.
Even though the cytosol is mostly water, it has a semi-solid, Jello-like consistency because of the
many proteins suspended in it. The cytosol contains a rich broth of macromolecules and smaller
organic molecules, including glucose and other simple sugars, polysaccharides, amino acids,
nucleic acids, and fatty acids. Ions of sodium, potassium, calcium, and other elements are also
found in the cytosol. Many metabolic reactions, including protein synthesis, take place in this part
of the cell.
Introduction
Suppose that you have a very precious piece of information. Let’s imagine that this piece of
information is a blueprint. In fact, it’s not just a blueprint for a house, or a car, or even a top-secret
fighter jet. It’s a blueprint for an entire organism – you – and it not only specifies how to put you
together, but also provides the information that enables every cell in your body to keep
functioning from moment to moment.
Sounds important, right? You’d probably want to keep information this valuable in a secure spot,
perhaps in a protected vault where you can keep an eye on it. In fact, that’s exactly what
eukaryotic cells do with their genetic material, placing it in a membrane-enclosed repository called
the nucleus.
Eukaryotic DNA never leaves the nucleus; instead, it’s transcribed (copied) into RNA molecules,
which may then travel out of the nucleus. In the cytosol, some RNAs associate with structures
called ribosomes, where they direct synthesis of proteins. (Other RNAs play functional roles in the
cell, serving as structural components of the ribosome or regulating activity of genes.) Here, we’ll
look in a little more detail at the structure of the nucleus and ribosomes.
[Do all cells have just one nucleus?]
The nucleus
The nucleus (plural, nuclei) houses the cell’s genetic material, or DNA, and is also the site of
synthesis for ribosomes, the cellular machines that assemble proteins. Inside the nucleus,
chromatin (DNA wrapped around proteins, described further below) is stored in a gel-like
substance called nucleoplasm.
Enclosing the nucleoplasm is the nuclear envelope, which is made up of two layers of
membrane: an outer membrane and an inner membrane. Each of these membranes contains two
layers of phospholipids, arranged with their tails pointing inward (forming a phospholipid bilayer).
There’s a thin space between the two layers of the nuclear envelope, and this space is directly
connected to the interior of another membranous organelle, the endoplasmic reticulum.
Nuclear pores, small channels that span the nuclear envelope, let substances enter and exit the
nucleus. Each pore is lined by a set of proteins, called the nuclear pore complex, that control
what molecules can go in or out.
If you look at a microscope image of the nucleus, you may notice – depending on the type of stain
used to visualize the cell – that there’s a dark spot inside it. This darkly staining region is called
the nucleolus, and it’s the site in which new ribosomes are assembled.
Diagram of the parts of the
nucleus of a eukaryotic
cell.
Image credit: OpenStax
Biology.
How do you make a ribosome? Some chromosomes have sections of DNA that encode
ribosomal RNA, a type of structural RNA that combines with proteins to make the ribosome. In
the nucleolus, new ribosomal RNA combines with proteins to form the subunits of the ribosome.
The newly made subunits are transported out through the nuclear pores to the cytoplasm, where
they can do their job.
Some cell types have more than one nucleolus inside the nucleus. For instance, some mouse
cells have up to 666 nucleoli^11start superscript, 1, end superscript. Prokaryotes, which do not
have a nucleus, don't have nucleoli and build their ribosomes in the cytosol.
[How is the nucleolus related to cancer?]
^{2,3}start superscript, 2, comma, 3, end superscript
Chromosomes and DNA
Now that we have a sense of the structure of the nucleus, let’s have a closer look at the genetic
information stored inside it: the DNA. Most of an organism’s DNA is organized into one or
more chromosomes, each of which is a very long string or loop of DNA. A single chromosome
can carry many different genes.
In prokaryotes, DNA is typically organized into a single circular chromosome (a loop). In
eukaryotes, on the other hand, chromosomes are linear structures (strings). Every eukaryotic
species has a specific number of chromosomes in the nuclei of its body’s cells. For example, a
typical human body cell would have 464646 chromosomes, while a comparable fruit fly cell would
have 888.
Chromosomes are only visible as distinct structures when the cell is getting ready to divide. When
the cell is in the growth and maintenance phases of its life cycle, the chromosomes instead
resemble an unwound, jumbled bunch of threads. In this form, the DNA is accessible to the
enzymes that transcribe it into RNA, allowing the genetic information to be put to use
(expressed).
In both their loose and compact forms, the DNA strands of chromosomes are bound to structural
proteins, including a family of proteins called histones (see picture below). These DNA-associated
proteins organize the DNA and help it fit into the nucleus, and they also play a role in determining
which genes are active or inactive. The complex formed by DNA and its supporting structural
proteins is known as chromatin. You can learn more about DNA, chromatin, and chromosomes
in the DNA and chromosomes article.
Left: image of a chromosome, showing how it is made up of DNA wound around histones and
then arranged in loops and other higher-order structures. Right: false-colored and rearranged
micrograph of chromosomes.
Image credit:
OpenStax
Biology.
Image at right
is a
modification of
work by the
NIH; scale-bar
data from Matt
Russell.
To give you a
sense of just
how important DNA packing is, consider that the DNA in a typical human cell would be
about 222 meters long if it were extended in a straight line. All 222 meters of that DNA are
squeezed into a tiny nucleus with a diameter of just 0.0060.0060, point, 006 mm. That's a feat
"geometrically equivalent to packing 404040 km (242424 miles) of extremely fine thread into a
tennis ball" ^44start superscript, 4, end superscript!
Ribosomes
As mentioned above, ribosomes are the molecular machines responsible for protein synthesis. A
ribosome is made out of RNA and proteins, and each ribosome consists of two separate RNA-
protein complexes, known as the small and large subunits. The large subunit sits on top of the
small subunit, with an RNA template sandwiched between the two. (A ribosome looks a little like
a hamburger with a puffy bun on top, an RNA “patty” threading through it.)
In eukaryotes, ribosomes get their orders for protein synthesis from the nucleus, where portions
of DNA (genes) are transcribed to make messenger RNAs (mRNAs). An mRNA travels to the
ribosome, which uses the information it contains to build a protein with a specific amino acid
sequence. This process is called translation. Prokaryotes lack a nucleus, so their mRNAs are
transcribed in the cytoplasm and can be translated by ribosomes immediately.
Image of a ribosome with small and large subunits, with an mRNA bound and a polypeptide chain
being produced. A tRNA molecule
is shown binding to the mRNA and
bringing the appropriate amino acid
into position for addition to the
chain.
Image credit: OpenStax Biology.
Mendoza, Alyssa Marie G.
Eukaryotic ribosomes may be either free, meaning that they are floating around in the cytoplasm,
or bound, meaning that they are attached to the endoplasmic reticulum or the outside of the
nuclear envelope. (In the first diagram in this article, the red dots represent bound ribosomes;
endoplasmic reticulum with bound ribosomes is known as rough endoplasmic reticulum.)
Because protein synthesis is an essential function of all cells, ribosomes are found in practically
every cell type of multicellular organisms, as well as in prokaryotes such as bacteria. However,
eukaryotic cells that specialize in producing proteins have particularly large numbers of
ribosomes. For example, the pancreas is responsible for producing and secreting large amounts
of digestive enzymes, so the pancreatic cells that make these enzymes have an unusually high
number of ribosomes.
Final fun fact: in a testament to the importance of the ribosome, the 2009 Nobel Prize in
Chemistry was awarded to three researchers who mapped its structure and movements down to
the level of individual atoms using a technique called X-ray crystallography^55start superscript, 5,
end superscript.
Introduction
Let’s imagine you are a pancreatic cell. Your job is to secrete digestive enzymes, which travel into
the small intestine and help break down nutrients from food. In order to carry out this job, you
somehow have to get those enzymes shipped from their site of synthesis—inside the cell—to
their place of action—outside the cell.
How are you going to make this happen? After a moment of panic in which you consider calling
the postal service, you relax, having remembered: I have an endomembrane system!
What is the endomembrane system?
The endomembrane system (endo- = “within”) is a group of membranes and organelles in
eukaryotic cells that works together to modify, package, and transport lipids and proteins. It
includes a variety of organelles, such as the nuclear envelope and lysosomes, which you may
already know, and the endoplasmic reticulum and Golgi apparatus, which we will cover shortly.
Micrograph and diagram of the endoplasmic reticulum. Micrograph shows the rough ER as a
series of membrane folds surrounding the nucleus. Diagram gives a 3D representation of rough
ER and smooth ER along with the cell nucleus.
_Image credit: left, "The endomembrane system and proteins: Figure 2" by OpenStax College,
Biology (CC BY 3.0), modification of work by Lousia Howard; right, modification of "Animal cell
structure" by Mariana Ruiz, public domain_
Since the rough ER helps modify proteins that will be secreted from the cell, cells whose job is to
secrete large amounts of enzymes or other proteins, such as liver cells, have lots of rough ER.
Smooth ER
The smooth endoplasmic reticulum (smooth ER) is continuous with the rough ER but has few
or no ribosomes on its cytoplasmic surface. Functions of the smooth ER include:
 Synthesis of carbohydrates, lipids, and steroid hormones
 Detoxification of medications and poisons

Storage of calcium ions
In muscle cells, a special type of smooth ER called the sarcoplasmic reticulum is responsible for
storage of calcium ions that are needed to trigger the coordinated contractions of the muscle
cells.
There are also tiny "smooth" patches of ER found within the rough ER. These patches serve as
exit sites for vesicles budding off from the rough ER and are called transitional ER^11start
superscript, 1, end superscript.
carbohydrate group attached to the protein.
Image showing transport of a membrane protein from the rough ER through the Golgi to the
plasma membrane. The protein is initially modified by the addition of branching carbohydrate
chains in the rough ER; these chains are then trimmed back and replaced with other branching
chains in the Golgi apparatus. The protein, with its final set of carbohydrate chains, is then
transported to the plasma membrane in a transport vesicle. The vesicle fuses with the plasma
membrane, its lipids and protein cargo becoming part of the plasma membrane.
_Image modified from "The endomembrane system and proteins: Figure 1" by OpenStax College,
Biology (CC BY 3.0), modification of work by Magnus Manske_
Finally, the modified proteins are sorted (based on markers such as amino acid sequences and
chemical tags) and packaged into vesicles that bud from the trans face of the Golgi. Some of
these vesicles deliver their contents to other parts of the cell where they will be used, such as the
lysosome or vacuole. Others fuse with the plasma membrane, delivering membrane-anchored
proteins that function there and releasing secreted proteins outside the cell.
Cells that secrete many proteins—such as salivary gland cells that secrete digestive enzymes, or
cells of the immune system that secrete antibodies—have many Golgi stacks. In plant cells, the
Golgi apparatus also makes polysaccharides (long-chain carbohydrates), some of which are
incorporated into the cell wall.
Lysosomes
The lysosome is an organelle that contains digestive enzymes and acts as the organelle-
recycling facility of an animal cell. It breaks down old and unnecessary structures so their
molecules can be reused. Lysosomes are part of the endomembrane system, and some vesicles
that leave the Golgi are bound for the lysosome.

Although it's not technically inside the cell, the plasma membrane is also part of the
endomembrane system. As we'll see, the plasma membrane interacts with the other
endomembrane organelles, and it's the site where secreted proteins (like the pancreatic enzymes
in the intro) are exported. Important note: the endomembrane system does not include
mitochondria, chloroplasts, or peroxisomes.
Let's take a closer look at the different parts of the endomembrane system and how they function
in the shipping of proteins and lipids.
The endoplasmic reticulum
The endoplasmic reticulum (ER) plays a key role in the modification of proteins and the
synthesis of lipids. It consists of a network of membranous tubules and flattened sacs. The discs
and tubules of the ER are hollow, and the space inside is called the lumen.
The Golgi apparatus [Why don't lysosomes digest themselves?]
When vesicles bud off from the ER, where do they go? Before reaching their final destination, the ^{1,2}start superscript, 1, comma, 2, end superscript
lipids and proteins in the transport vesicles need to be sorted, packaged, and tagged so that they
wind up in the right place. This sorting, tagging, packaging, and distribution takes place in
the Golgi apparatus(Golgi body), an organelle made up of flattened discs of membrane. Lysosomes can also digest foreign particles that are brought into the cell from outside. As an
example, let's consider a class of white blood cells called macrophages, which are part of the
human immune system. In a process known as phagocytosis, a section of the macrophage’s
plasma membrane invaginates—folds inward—to engulf a pathogen, as shown below.
Micrograph of the Golgi apparatus showing a series of flattened membrane discs in cross-section
_Image credit: "The endomembrane system and proteins: Figure 3" by OpenStax College,
Biology (CC BY 3.0), modification of work by Lousia Howard_

Diagram of
Rough ER phagocytosis, in
which the
phagosome
The rough endoplasmic reticulum (rough ER) gets its name from the bumpy ribosomes generated by
attached to its cytoplasmic surface. As these ribosomes make proteins, they feed the newly engulfment of a
forming protein chains into the lumen. Some are transferred fully into the ER and float inside, particle fuses
while others are anchored in the membrane. with a
lysosome,
Inside the ER, the proteins fold and undergo modifications, such as the addition of carbohydrate allowing
side chains. These modified proteins will be incorporated into cellular membranes—the digestion of the
membrane of the ER or those of other organelles—or secreted from the cell. particle.

The receiving side of the _Image credit:

If the modified proteins are not destined to stay in the ER, they will be packaged into vesicles, or Golgi apparatus is called
small spheres of membrane that are used for transport, and shipped to the Golgi apparatus. The
the cis face and the opposite
rough ER also makes phospholipids for other cellular membranes, which are transported when side is called the trans face.
the vesicle forms.
Transport vesicles from the
ER travel to the cisface, fuse
with it, and empty their
contents into the lumen of
the Golgi apparatus.
modified from
"The endomembrane system and proteins: Figure
4" by OpenStax College, Biology (CC BY 3.0)_
The invaginated section, with the pathogen inside, pinches off from the plasma membrane to form
a structure called a phagosome. The phagosome then fuses with a lysosome, forming a
combined compartment where digestive enzymes destroy the pathogen.

As proteins and lipids travel Vacuoles

through the Golgi, they
undergo further
modifications. Short chains
of sugar molecules might be
added or removed, or
phosphate groups attached
as tags. Carbohydrate
processing is shown in the
diagram as the gain and loss
of branches on the purple
Plants cells are unique because they have a lysosome-like organelle called the vacuole. The
large central vacuole stores water and wastes, isolates hazardous materials, and has enzymes
that can break down macromolecules and cellular components, like those of a lysosome.^33start
superscript, 3, end superscript Plant vacuoles also function in water balance and may be used to
store compounds such as toxins and pigments (colored particles).^44start superscript, 4, end
superscript
 Mendoza, Alyssa Marie G.
Lysosomes vs. peroxisomes
One point that can be confusing is the difference between lysosomes and peroxisomes. Both
types of organelles are involved in breaking down molecules and neutralizing hazards to the cell.
Also, both usually show up as small, round blobs in diagrams.
However, the peroxisome is a different organelle with its own unique properties and role in the
cell. It houses enzymes involved in oxidation reactions, which produce hydrogen peroxide (\text
H_2 \text O_2H2O2H, start subscript, 2, end subscript, O, start subscript, 2, end subscript) as a
by-product. The enzymes break down fatty acids and amino acids, and they also detoxify some
substances that enter the body. For example, alcohol is detoxified by peroxisomes found in liver
cells.
Importantly, peroxisomes—unlike lysosomes—are not part of the endomembrane system. That
means they don't receive vesicles from the Golgi apparatus. You can learn more about how
proteins are shipped to the peroxisome in the article on protein targeting.
Key points:
 Mitochondria are the "powerhouses" of the cell, breaking down fuel molecules and capturing
energy in cellular respiration.
 Chloroplasts are found in plants and algae. They're responsible for capturing light energy to
make sugars in photosynthesis.
 Mitochondria and chloroplasts likely began as bacteria that were engulfed by larger cells
(the endosymbiont theory).
Introduction
You may know that your body is made up of cells (trillions and trillions of them). You may also
know that the reason you need to eat food—such as veggies—is so that you have the energy to
do things like play sports, study, walk, and even breathe.
But what exactly happens in your body to turn the food energy stored in broccoli into a form that
your body can use? And how does energy end up stored in the broccoli to begin with, anyway?
The answers to these questions have a lot to do with two important organelles: mitochondria and
chloroplasts.
 Chloroplasts are organelles found in the broccoli's cells, along with those of other plants and
algae. They capture light energy and store it as fuel molecules in the plant's tissues.
 Mitochondria are found inside of your cells, along with the cells of plants. They convert the energy
stored in molecules from the broccoli (or other fuel molecules) into a form the cell can use.
Let's take a closer look at these two very important organelles.
Chloroplasts
Chloroplasts are found only in plants and photosynthetic algae. (Humans and other animals do
not have chloroplasts.) The chloroplast's job is to carry out a process called photosynthesis.
In photosynthesis, light energy is collected and used to build sugars from carbon dioxide. The
sugars produced in photosynthesis may be used by the plant cell, or may be consumed by
animals that eat the plant, such as humans. The energy contained in these sugars is harvested
through a process called cellular respiration, which happens in the mitochondria of both plant and
animal cells.
Chloroplasts are disc-shaped organelles found in the cytosol of a cell. They have outer and inner
membranes with an intermembrane space between them. If you passed through the two layers of
membrane and reached the space in the center, you’d find that it contained membrane discs
known as thylakoids, arranged in interconnected stacks called grana (singular, granum).
Diagram of a chloroplast, showing the
outer membrane, inner membrane,
intermembrane space, stroma, and
thylakoids arranged in stacks called
grana.
_Image modified from "Chloroplast mini,"
by Kelvin Ma (CC BY 3.0)._
The membrane of a thylakoid disc
contains light-harvesting complexes
that include chlorophyll, a pigment that
gives plants their green color. Thylakoid
discs are hollow, and the space inside a disc
is called the thylakoid space or lumen, while the fluid-filled space surrounding the thylakoids is work?]
called the stroma.
\text H^+H, start superscript, plus, end superscript
You can learn more about chloroplasts, chlorophyll, and photosynthesis in
the photosynthesis topic section.
Electrons from fuel molecules, such as the sugar glucose, are stripped off in reactions that take
place in the cytosol and in the mitochondrial matrix. These electrons are captured by special
Mitochondria molecules called electron carriers and deposited into the electron transport, a series of proteins
embedded in the inner mitochondrial membrane. As the electrons move down the transport chain,
energy is released and used to pump protons (\text H^+H+H, start superscript, plus, end
Mitochondria (singular, mitochondrion) are often called the powerhouses or energy factories of superscript) out of the matrix and into the intermembrane space. As protons flow back down their
the cell. Their job is to make a steady supply of adenosine triphosphate (ATP), the cell’s main gradient and into the matrix, they pass through an enzyme called ATP synthase, which harnesses
energy-carrying molecule. The process of making ATP using chemical energy from fuels such as the flow of protons to generate ATP from ADP and Pi.
sugars is called cellular respiration, and many of its steps happen inside the mitochondria.
Although mitochondria are found in most human cell types (as well as most cell types in other
The mitochondria are suspended in the jelly-like cytosol of the cell. They are oval-shaped and animals and plants), their numbers vary depending on the role of the cell and its energy needs.
have two membranes: an outer one, surrounding the whole organelle, and an inner one, with For instance, muscle cells typically have high energy needs and large numbers of mitochondria,
many inward protrusions called cristae that increase surface area. while red blood cells, which are highly specialized for oxygen transport, have no mitochondria at
all.^33start superscript, 3, end superscript
Where did these organelles come from?
Both mitochondria and chloroplasts contain their own DNA and ribosomes. Why would these
organelles need DNA and ribosomes, when there is DNA in the nucleus and ribosomes in the
Electron micrograph of a mitochondrion,
showing matrix, cristae, outer membrane, and
cytosol?
inner membrane.
Strong evidence points to endosymbiosis as the answer to the puzzle. Symbiosis is a
relationship in which organisms from two separate species live in a close, dependent relationship.
_Image credits: upper image, "Eukaryotic cells:
Endosymbiosis (endo- = “within”) is a specific type of symbiosis where one organism lives inside
Figure 7," by OpenStax College, Biology (CC BY
the other.
3.0). Modification of work by Matthew Britton; scale-
bar data from Matt Russell. Lower image: modification
of "Mitochondrion mini," by Kelvin Ma (public
domain)._
1. The first endosymbiotic event occurred: The ancestral eukaryote consumed aerobic bacteria that
Cristae were once thought to be broad, wavy evolved into mitochondria.
folds, but as Sal discusses in his mitochondria 2. In a second endosymbiotic event, the early eukaryote consumed photosynthetic bacteria that
video, they're now understood to be more like long evolved into chloroplasts."
caverns.^11start superscript, 1, end superscriptHere is a 3D reconstruction of a slice cut from a _Image modified from
mitochondrion: "Eukaryotic origins: Figure 4,"
by OpenStax College, Biology,
(CC BY 4.0)._
Bacteria, mitochondria,
and chloroplasts are
similar in size. Bacteria
also have DNA and
ribosomes similar
to those of
mitochondria
and
chloroplasts.^44start superscript, 4, end superscript Based on this and other evidence, scientists
think host cells and bacteria formed endosymbiotic relationships long ago, when individual host
cells took in aerobic (oxygen-using) and photosynthetic bacteria but did not destroy them.
Through millions of years of evolution, the aerobic bacteria became mitochondria and the
photosynthetic bacteria became chloroplasts.
Introduction
Image What would happen if someone snuck in during the night and stole your skeleton? Just to be
credit: "MitochondrionCAM," by Carmann (public domain).^22start superscript, 2, end superscript
clear, that’s not very likely to happen, biologically speaking. But if it did somehow happen, the
loss of your skeleton would cause your body to lose much of its structure. Your external shape
would change, some of your internal organs might start moving out of place, and you would
The space between the membranes is called the intermembrane space, and the compartment
probably find it very difficult to walk, talk, or move.
enclosed by the inner membrane is called the mitochondrial matrix. The matrix contains
mitochondrial DNA and ribosomes. We'll talk shortly about why mitochondria (and chloroplasts)
have their own DNA and ribosomes. Interestingly enough, the same is true for a cell. We often think about cells as soft, unstructured
blobs. But in reality, they are highly structured in much the same way as our own bodies. They
have a network of filaments known as the cytoskeleton (literally, “cell skeleton”), which not only
The multi-compartment structure of the mitochondrion may seem complicated to us. That's true,
supports the plasma membrane and gives the cell an overall shape, but also aids in the correct
but it turns out to be very useful for cellular respiration, allowing reactions to be kept separate and positioning of organelles, provides tracks for the transport of vesicles, and (in many cell types)
different concentrations of molecules to be maintained in different "rooms." allows the cell to move.
In eukaryotes, there are three types of protein fibers in the cytoskeleton: microfilaments,
intermediate filaments, and microtubules. Here, we'll examine each type of filament, as well as
[How
some specialized structures related to the cytoskeleton.
does that
Microfilaments
Mendoza, Alyssa Marie G.
Of the three types of protein fibers in the cytoskeleton, microfilaments are the narrowest. They
have a diameter of about 7 nm and are made up of many linked monomers of a protein called
actin, combined in a structure that resembles a double helix. Because they are made of actin
monomers, microfilaments are also known as actin filaments. Actin filaments have directionality,
meaning that they have two structurally different ends.
Actin filament, a double helix-like structure made out of actin monomers
(subunits).
Image modified from OpenStax Biology.
Actin filaments have a number of important roles in the cell. For one, they
serve as tracks for the movement of a motor protein called myosin, which
can also form filaments. Because of its relationship to myosin, actin is
involved in many cellular events requiring motion.
For instance, in animal cell division, a ring made of actin and myosin
pinches the cell apart to generate two new daughter cells. Actin and myosin
are also plentiful in muscle cells, where they form organized structures of
overlapping filaments called sarcomeres. When the actin and myosin filaments of a sarcomere
slide past each other in concert, your muscles contract.
Actin filaments may also serve as highways inside the cell for the transport of cargoes, including
protein-containing vesicles and even organelles. These cargoes are carried by individual myosin
motors, which "walk" along actin filament bundles^11start superscript, 1, end superscript.
Actin filaments can assemble and disassemble quickly, and this property allows them to play an
important role in cell motility (movement), such as the crawling of a white blood cell in your
immune system.
Finally, actin filaments play key structural roles in the cell. In most animal cells, a network of actin
filaments is found in the region of cytoplasm at the very edge of the cell. This network, which is
linked to the plasma membrane by special connector proteins, gives the cell shape and
structure^{2,3}2,3start superscript, 2, comma, 3, end superscript.
Intermediate filaments
Intermediate filaments are a type of cytoskeletal element made of multiple strands of fibrous
proteins wound together. As their name suggests, intermediate filaments have an average
diameter of 8 to 10 nm, in between that of microfilaments and microtubules (discussed below).
Intermediate filaments in a cell, forming a networks that holds the nucleus and other organelles in
place.Image credit:
"The cytoplasm and cellular organelles," by OpenStax College
(CC BY 3.0).
Intermediate filaments come in a number of different varieties,
each one made up of a different type of protein. One protein
that forms intermediate filaments is keratin, a fibrous protein
found in hair, nails, and skin. For instance, you may have seen
shampoo ads that claim to smooth the keratin in your hair!
Unlike actin filaments, which can grow and disassemble
quickly, intermediate filaments are more permanent and play
an essentially structural role in the cell. They are specialized to
bear tension, and their jobs include maintaining the shape of
the cell and anchoring the nucleus and other organelles in place.
Microtubules
Despite the “micro” in their name, microtubules are the largest of the three types of cytoskeletal
fibers, with a diameter of about 25 nm. A microtubule is made up of tubulin proteins arranged to
form a hollow, straw-like tube, and each tubulin protein consists of two subunits, α-tubulin and β-
tubulin.
Microtubules, like actin filaments, are dynamic structures: they can grow and shrink quickly by the
addition or removal of tubulin
proteins. Also similar to actin
filaments, microtubules have
directionality, meaning that
they have two ends that are
structurally different from
one another. In a cell,
microtubules play an important structural role, helping the cell resist compression forces.
Left: 3D model of a microtubule, showing that it is a hollow cylinder of proteins. Right: Cartoon
diagram of a microtubule, showing that it is made of two different types of subunits (alpha and
beta). The subunits form dimers, and the dimers are connected in a spiral pattern to form the
hollow tube of the microtubule.
Image credit: OpenStax Biology.
In addition to providing structural support, microtubules play a variety of more specialized roles in
a cell. For instance, they provide tracks for motor proteins called kinesins and dyneins, which
transport vesicles and other cargoes around the interior of the cell^44start superscript, 4, end
superscript. During cell division, microtubules assemble into a structure called the spindle, which
pulls the chromosomes apart.
Flagella, cilia, and centrosomes
Microtubules are also key components of three more specialized eukaryotic cell structures:
flagella, cilia and centrosomes. You may remember that our friends the prokaryotes also have
structures have flagella, which they use to move. Don't get confused—the eukaryotic flagella
we're about to discuss have pretty much the same role, but a very different structure.
Flagella (singular, flagellum) are long, hair-like structures that extend from the cell surface and
are used to move an entire cell, such as a sperm. If a cell has any flagella, it usually has one or
just a few. Motile cilia (singular, cilium) are similar, but are shorter and usually appear in large
numbers on the cell surface. When cells with motile cilia form tissues, the beating helps move
materials across the surface of the tissue. For example, the cilia of cells in your upper respiratory
system help move dust and particles out towards your nostrils.
Despite their difference in length and number, flagella and motile cilia share a common structural
pattern. In most flagella and motile cilia, there are 9 pairs of microtubules arranged in a circle,
along with an additional two microtubules in the center of the ring. This arrangement is called a 9
+ 2 array. You can see the 9 + 2 array in the electron microscopy image at left, which shows two
flagella in cross-section.
Upper: Transmission electron micrograph of flagella in cross-
section, showing the 9+2 microtubule array organization.
Lower: Cartoon diagram of a motile cililum, showing the singlet
microtubules in the center, the outer doublet microtubules
arranged in a circle around the singlet microtubules, and the
dyneins attached to the doublet microtubules. The whole structure
is surrounded by plasma membrane. At the base of the cilium lies
a basal body, which is also made up of microtubules.
_Image credits: upper panel, "The cytoskeleton: Figure 5," by
OpenStax College, Biology (CC BY 3.0). Modification of work by
Dartmouth Electron Microscope Facility, Dartmouth College; scale-
bar data from Matt Russell. Lower panel, modification of
"Eukaryotic cilium diagram," by Mariana Ruiz Villareal (public domain)._
In flagella and motile cilia, motor proteins called dyneins move along the microtubules, generating
a force that causes the flagellum or cilium to beat. The structural connections between the
microtubule pairs and the coordination of dynein movement allow the activity of the motors to
produce a pattern of regular beating^{5,6}5,6start superscript, 5, comma, 6, end superscript.
You may notice another feature in the diagram above: the cilium or flagellum has a basal
body located at its base. The basal body is made of microtubules and plays a key role in
assembly of the cilium or flagellum. Once the structure has been assembled, it also regulates
which proteins can enter or exit^77start superscript, 7, end superscript.
The basal body is actually just a modified centriole^77start superscript, 7, end superscript.
A centriole is a cylinder of nine triplets of microtubules, held together by supporting proteins.
Centrioles are best known for their role in centrosomes, structures that act as microtubule
organizing centers in animal cells. A
centrosome consists of two centrioles
oriented at right angles to each other,
surrounded by a mass of "pericentriolar
material," which provides anchoring sites
for microtubules^88start superscript, 8,
end superscript.
Image of a centrosome. The centrosome
contains two centrioles positioned at right
angles to each other.
Image credit: modification of "Centriole," by Kelvinsong (CC BY 3.0)
The centrosome is duplicated before a cell divides, and the paired centrosomes seem to play a
role in organizing the microtubules that separate chromosomes during cell division. However, the
exact function of the centrioles in this process still isn’t clear. Cells with their centrosome removed
can still divide, and plant cells, which lack centrosomes, divide just fine.
Introduction
We’ve spent a lot of time looking at what’s inside a cell. What, then, is on the outside? It depends
a lot on what kind of cell you’re looking at.
Plants and fungi have a tough cell wall for protection and support, while animal cells can secrete
materials into their surroundings to form a meshwork of macromolecules called the extracellular
matrix. Here, we’ll look in more detail at these external structures and the roles they play in
different cell types.
Extracellular matrix of animal cells
Most animal cells release materials into the extracellular space, creating a complex meshwork of
proteins and carbohydrates called the extracellular matrix (ECM). A major component of the
extracellular matrix is the protein collagen. Collagen proteins are modified with carbohydrates,
and once they're released from the cell, they assemble into long fibers called collagen
fibrils^{1}1start superscript, 1, end superscript.
Collagen plays a key role in giving tissues strength and structural integrity. Human genetic
disorders that affect collagen, such as Ehlers-Danlos syndrome, result in fragile tissues that
stretch and tear too easily^22start superscript, 2, end superscript.
In the extracellular matrix, collagen fibers are interwoven with a class of carbohydrate-
bearing proteoglycans, which may be attached to a long polysaccharide backbone as shown in
the picture below. The extracellular matrix also contains many other types of proteins and
carbohydrates.
[More about proteoglycans]
^1start superscript, 1, end superscript
Diagram showing the extracellular
matrix and its connections to the cell. A
network of collagen fibers and
proteoglycans is found outside of the
cell. Collagen connects to integrin
proteins in the plasma membrane via
fibronectin. On the inside of the cell, the
integrins link up to the microfilaments of
the cytoskeleton.
Image credit: OpenStax Biology.
The extracellular matrix is directly
connected to the cells it surrounds.
Some of the key connectors are
proteins called integrins, which are
embedded in the plasma membrane.
Proteins in the extracellular matrix, like
the fibronectinmolecules shown in
green in the diagram above, can act as
bridges between integrins and other
extracellular matrix proteins such as collagen. On the inner side of the membrane, the integrins
are linked to the cytoskeleton.
Integrins anchor the cell to the extracellular matrix. In addition, they help it sense its environment.
They can detect both chemical and mechanical cues from the extracellular matrix and trigger
signaling pathways in response^{4,5}4,5start superscript, 4, comma, 5, end superscript.
Blood clotting provides another example of communication between cells and the extracellular
matrix. When the cells lining a blood vessel are damaged, they display a protein receptor called
tissue factor. When tissue factor binds to a molecule present in the extracellular matrix, it triggers
a range of responses that reduce blood loss. For instance, it causes platelets to stick to the wall
of the damaged blood vessel and stimulates them to produce clotting factors.
The cell wall
Though plants don't make collagen, they have their own type of supportive extracellular structure:
the cell wall. The cell wall is a rigid covering that surrounds the cell, protecting it and giving it
Mendoza, Alyssa Marie G.
support and shape. Have you ever noticed that when you bite into a raw vegetable, like celery, it
crunches? A big part of that crunch is the rigidity of celery’s cell walls.
Fungi also have cell walls, as do some protists (a group of mostly unicellular eukaryotes) and
most prokaryotes—though I don't recommend biting into any of those to see if they crunch. Like
the animal extracellular matrix, the plant cell wall is made up of molecules secreted by the cell.
The major organic molecule of the plant cell wall is cellulose, a polysaccharide composed of
glucose units. Cellulose assembles into fibers called microfibrils, as shown in the diagram below.
Functionally, gap junctions in animal cells are a lot like plasmodesmata in plant cells: they are Image credit: OpenStax Biology. Modification of work by Mariana Ruiz Villareal.
channels between neighboring cells that allow for the transport of ions, water, and other
substances^33start superscript, 3, end superscript. Structurally, however, gap junctions and
plasmodesmata are quite different. Desmosomes pin adjacent cells together, ensuring that cells in organs and tissues that stretch,
such as skin and cardiac muscle, remain connected in an unbroken sheet.
In vertebrates, gap junctions develop when a set of six membrane proteins
called connexins form an elongated, donut-like structure called a connexon. When the pores, or
“doughnut holes,” of connexons in adjacent animal cells align, a channel forms between the cells.
(Invertebrates also form gap junctions in a similar way, but use a different set of proteins called
innexins.)^44start superscript, 4, end superscript

Image of the plant cell wall, showing the network of cellulose microfibrils and pectins (with pectins
being particularly abundant in the middle lamella).

Image credit: "Plant cell wall diagram" by Mariana Ruiz Villareal, public domain

Most plant cell walls contain a

variety of different
polysaccharides and proteins. In
addition to cellulose, other
polysaccharides commonly found
in the plant cell wall include
hemicellulose and pectin, shown
in the diagram above. The middle
lamella, shown along the top of
the diagram, is a sticky layer that
helps hold the cell walls of
adjacent plant cells
together^{6,7}6,7start superscript,
6, comma, 7, end superscript.
Image of the plasma membranes of two cells held
together by gap junctions. Where two connexons from
the different cells meet, they can form a channel
leading from one cell into the next.
Image credit: OpenStax Biology. Modification of work
by Mariana Ruiz Villareal.
Gap junctions are particularly important in cardiac muscle: the electrical signal to contract
spreads rapidly between heart muscle cells as ions pass through gap junctions, allowing the cells
to contract in tandem.

Introduction Tight junctions

If you were building a building, what kinds of connections might you want to put between the
rooms? In some cases, you’d want people to be able to walk from one room to another, in which
case you’d put in a door. In other cases, you’d want to hold two adjacent walls firmly together, in
which case you might put in some strong bolts. And in still other cases, you might need to ensure
that the walls were sealed very tightly together – for instance, to prevent water from dripping
between them.
As it turns out, cells face the same questions when they’re arranged in a tissue next to other cells.
Should they put in doors that connect them directly to their neighbors? Do they need to spot-weld
themselves to their neighbors to make a strong layer, or perhaps even form tight seals to prevent
water from passing through the tissue? Junctions serving all of these functions can be found in
cells of different types, and here, we’ll look at each of them in turn.
Plasmodesmata
Not all junctions between cells produce cytoplasmic connections. Instead, tight junctions create
a watertight seal between two adjacent animal cells.
At the site of a tight junction, cells are held tightly against each other by many individual groups of
tight junction proteins called claudins, each of which interacts with a partner group on the
opposite cell membrane. The groups are arranged into strands that form a branching network,
with larger numbers of strands making for a tighter seal^55start superscript, 5, end superscript.
Image of the membranes of two cells held
together by tight junctions. The tight
junctions are like rivets, and they are
arranged in multiple strands that form
lines and triangles.

Image credit: OpenStax Biology.

Plant cells, surrounded as they are by cell walls, don’t contact one another through wide stretches
of plasma membrane the way animal cells can. However, they do have specialized junctions
called plasmodesmata (singular, plasmodesma), places where a hole is punched in the cell wall
to allow direct cytoplasmic exchange between two cells.
Image of two cells connected by a plasmodesma, showing how materials can travel from the
cytoplasm of one cell to
the next via the
plasmodesma.
Modification of work by Mariana Ruiz
Villareal.
The purpose of tight junctions is to keep
liquid from escaping between cells,
allowing a layer of cells (for instance,
those lining an organ) to act as an
impermeable barrier. For example, the tight junctions between the epithelial cells lining your
bladder prevent urine from leaking out into the extracellular space.

Image credit: OpenStax Desmosomes

Biology.

Animal cells may also contain junctions called desmosomes, which act like spot welds between
Plasmodesmata are lined adjacent epithelial cells. A desmosome involves a complex of proteins. Some of these proteins
with plasma membrane extend across the membrane, while others anchor the junction within the cell.
that is continuous with
the membranes of the
two cells. Each Cadherins, specialized
plasmodesma has a adhesion proteins, are found on
thread of cytoplasm extending through it, containing an even thinner thread of endoplasmic the membranes of both cells and
reticulum (not shown in the diagram above). interact in the space between
them, holding the membranes
together. Inside the cell, the
Molecules below a certain size (the size exclusion limit) move freely through the plasmodesmal cadherins attach to a structure
channel by passive diffusion. The size exclusion limit varies among plants, and even among cell called the cytoplasmic plaque
types within a plant. Plasmodesmata may selectively dilate (expand) to allow the passage of (red in the image at right), which
certain large molecules, such as proteins, although this process is poorly connects to the intermediate
understood^{1,2}1,2start superscript, 1, comma, 2, end superscript. filaments and helps anchor the
junction.
Gap junctions