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2 Membranes and
cell organelles
This chapter is designed to enable students to: • distinguish the different ways in which
• understand the extent of the plasma biomolecules enter or leave cells
membrane in forming a series of membranous • develop their knowledge and understanding of
channels for the packaging and transport of connections between cells
biomolecules throughout eukaryote cells • extend their understanding of apoptosis.
• enhance their knowledge and understanding
of the structure and function of cell
organelles
Figure 2.1 This image shows a transverse section of a mouse eukaryotic cells and all share the characteristic of an internal
tail. Look at the incredible range of different kinds of cells structure of membranous chambers called organelles. In this
present: cartilage, connective tissue, nerve, muscle, epithelial chapter we consider the structure and function of organelles.
cells and others. The nucleus of each cell contains the same We also consider the transport of material within cells and the
DNA. Although some proteins are made by all cells, others are passage of material across plasma membranes.
32 NATURE
different OF BIOLOGY
and give BOOKof1 cell its uniqueness. These are
each kind
Life or death for a cell?
Groups of similar cells form tissues and groups of tissues come together to form
organs. The death of cells is a natural feature of healthy tissue. This ‘programmed
cell death’ was first noted in 1972 by Andrew Wyllie and is called apoptosis
(from Greek, meaning ‘shedding of leaves in autumn’).
In fully formed tissue, cell death and cell reproduction are generally balanced.
If this balance is not regulated, an uncontrolled increase in cells can occur and a
tumour develops. If a tumour continues to grow and invades healthy tissue, it is
said to be malignant. A malignant tumour is called cancer. Too little apoptosis can
lead to cancer and too much can cause degenerative diseases such as Alzheimer
disease.
Pancreas (2%)
Cervix (4%)
Ovary (4%)
Colo-rectal (15%)
Uterus (4%)
Cancer is the second highest cause of death after heart disease in Australia and
Figure 2.3 Mouse fibroblast cells. breast cancer is the most common cause affecting adult females (see figure 2.2).
Fibroblasts are common in areolar Although there has been improvement in the treatment of cancers in recent years,
tissue, a connective tissue found 30 per cent of women diagnosed with breast cancer will die from it. Researchers
below the skin, around blood vessels in the Cancer Division at the Walter and Eliza Hall Institute of Medical Research
and nerves, and filling the spaces (WEHI) in Melbourne are investigating how breast cancer develops. This involves
between organs. When images of identifying regulator proteins within cells and investigating the interactions of
Bim proteins (stained red) and Bcl-2 these proteins that ultimately decide whether a cell lives or dies. Special stains,
proteins (stained green) within a cell such as those used on the cells in figure 2.3, assist in pinpointing the positions
are superimposed, a yellow colour
of regulator proteins within cells. Other experiments are aimed at establishing
results. This indicates that the two
proteins, which are both associated
the physiological role of these proteins. If we have better information about
with programmed cell death the control and development of cancers, there is an increased chance that better
(apoptosis), are bound to the same treatments can be developed.
membranes within the cytoplasm of Read about Sue Macaulay’s work as a radiographer with St Vincent’s Breast-
the cell. Screen service, on page 36.
10 µm 10 µm 10 µm
QUICK-CHECK
1 Why are cells known as the basic building blocks of living organisms?
2 How might an examination of cells help diagnose disease?
APOPTOSIS
Apoptosis, or programmed cell death, is self-destruction Signals from outside a cell —
by cells for the good of the whole organism. What is the death receptor pathway
the difference between this type of cell death and the
type that we call necrosis? Necrosis occurs if a cell is Why would a perfectly healthy cell receive a message
seriously damaged by some mechanical or chemical to self-destruct? There are different reasons. The signal
trauma and this causes general damage to the plasma that a cell may receive could be:
membrane of the cell. the plasma membrane can no • You haven’t developed fully. This occurs in the
longer control what enters or leaves the cell, the cell embryonic brain when billions of cells are formed
swells then bursts and the contents spread out over but some fail to be incorporated accurately into the
nearby cells, causing inflammation of those tissues. brain network. These ‘stray’ cells die by apoptosis.
In apoptosis, cells respond to signals. There are two • There are more of you than are needed. It ‘costs’ an
main pathways of signals that initiate apoptosis: the organism energy and materials to keep unneeded
mitochondrial pathway and the death receptor pathway. cells alive. Some immune system cells are produced
in larger numbers than required. These excess cells
Signals from inside a cell — the die by apoptosis.
mitochondrial pathway
(a)
If serious damage occurs inside a cell, for example,
severe DNA damage or malfunction of an oxidative
enzyme, proteins on the surface of mitochondria are
activated and the mitochondrial membrane breaks.
This starts a series of events in the cell, including the
action of caspases (special enzymes that cleave specific
proteins at the amino acid aspartite) which enter the
nuclear pores and break DNA into small pieces. Events
after this are similar to those described below for signals
from outside a cell.
Another situation in which a cell may initiate death
itself is if a cell is infected with a virus. The cell identi-
fies the infection and kills itself before the virus has had
time to replicate and spread to other cells.
• Phagocytosis of parts
• Cytokines secreted
• Components recycled
• Organelles recycled
QUICK-CHECK
3 List three possible death signals a cell might receive to initiate
apoptosis.
1 mm
100 µm
10 µm
1 µm
0.1 µm
0.01 µm
0.001 µm
0.0001 µm
Organelles are held in place by a network of fine protein filaments within a
cell, collectively known as the cytoskeleton (see page 52). Prokaryotic cells such
as bacteria lack these internal membranes.
In the following sections, we examine the plasma membrane and cell organelles
of eukaryotes and discuss their functions.
Table 2.1 Relative volumes of the major compartments within a liver cell
Intracellular compartment Percentage of total cell volume
Cytoplasm = cytosol + organelles Cytosol 54
except the nucleus
Protoplasm = cytosol + all Mitochondria 22
organelles
Rough endoplasmic reticulum 9
Smooth endoplasmic reticulum 6
Nucleus 6
Lysosomes, peroxisomes, endosomes 3
Table 2.2 Relative amounts of membrane associated with some of the organelles in two
different kinds of cells
Nucleus
Mitochondrion
Ribosome
Endoplasmic
reticulum
Lysosome
Figure 2.8 The outer plasma
membrane of a typical animal Centriole
cell contains a network of inner
membranes that create smaller Peroxisome
compartments within the cell, known
as organelles. We will discuss the Protein
plasma membrane and the organelles microtubule
labelled in red in the following Golgi apparatus
sections.
Vesicle
Outside cell
Glycolipid
Phospholipid
bilayer
Membrane
glycoprotein
Figure 2.9 The plasma membrane
of all cells has the same basic
Cytoplasm
structure. Note the phospholipid
bilayer. Proteins penetrate into or
through the phospholipid bilayer
and carbohydrate chains bond to
many of these. A few carbohydrate
chains bond directly to the outer Nucleus
phospholipid layer. Note the pores
in the nuclear membrane.
Free passage
Diffusion is the net movement of a substance, typically in a solution, from a
ODD FACT region of high concentration of the substance to a region of low concentration
(see figure 2.10a). The process of diffusion does not require energy.
One special case At all times, molecules are in random movement. If a substance is more con-
of diffusion is known as centrated outside the cell than inside, molecules move from outside to inside the
osmosis. The process of osmosis cell. Diffusion stops at the stage when the concentration of substance X is equal
occurs when a net movement on the two sides of the membrane.
of water molecules occurs by Substances that can dissolve readily in water are termed hydrophilic, or
diffusion across a cell membrane
‘water-loving’. Some substances that have low water solubility or do not dissolve
either into or out of a cell.
in water are able to dissolve in or mix uniformly with lipid. These substances
are termed lipophilic (sometimes called hydrophobic). Examples of lipophilic
substances include alcohol and ether. Lipophilic substances can cross plasma
membrane boundaries readily.
Channel mediated
Some substances that are unable to carry out simple diffusion through the phos-
pholipid bilayer gain free passage across a membrane with the assistance of
protein channels (see figure 2.10b). Molecules move from a high concentration
to a low concentration without requiring energy.
Carrier mediated
Sometimes a protein channel alone is insufficient and a carrier molecule is
required to move molecules down the concentration gradient through a protein
channel (see figure 2.10c). When a specific carrier molecule is required, this kind
of movement is also called facilitated diffusion.
Movement of substances by facilitated diffusion mainly involves substances
that cannot diffuse across the plasma membrane by dissolving in the phospholipid
bilayer of the membrane. For example, the movement of glucose molecules across
the plasma membrane of red blood cells involves a specific carrier molecule.
All three methods of passive transport (figure 2.10a–c) result in molecules
moving from a region of high concentration to a region of low concentration
without the expenditure of energy.
Outside Outside
cell cell
Phospholipid
bilayer
Concentration
gradient in all
Energy the cases shown
Inside
cell
Inside
cell
Figure 2.10 Transport of molecules across membranes: (a–c) Three ways in which molecules move from a region
of high concentration, across a plasma membrane, to a region of low concentration without the expenditure of energy.
(d) Movement of molecules from a region of low concentration across a plasma membrane to a region of high concentration
requires the expenditure of energy. Note the movement of molecules against the concentration gradient.
Lipid
bilayer
Cytosol Cytosol
Lysosome
KEY IDEAS
• Each eukaryotic cell contains many membranous structures, called
organelles, suspended in the cytosol.
• Every living cell has a plasma membrane boundary.
• There are several different ways in which materials cross plasma
membranes to enter cells.
• Cell walls lie outside the plasma membrane of plant, fungal and
prokaryotic cells.
QUICK-CHECK
Nucleus
4 Make a labelled sketch of a typical plasma membrane.
5 List the different ways in which materials cross plasma membranes.
For each way, indicate whether or not it is energy-requiring.
6 Many plant cells have secondary cell walls as well as primary. Of what
advantage is this to a plant?
KEY IDEAS
• Nucleoli contain the nucleic acid RNA.
• The nucleus contains the nucleic acid DNA, which is the genetic
material within a cell.
• The nucleus of eukaryote cells is enclosed within a nuclear envelope.
QUICK-CHECK
7 State whether the following are true or false and briefly explain your
answer.
a A nucleus from a plant cell would be expected to have a double
nuclear membrane.
b Chromosomes are always visible in a eukaryotic cell.
8 Suggest why the nucleus is sometimes called the ‘control centre’ of a
cell.
NH2
C N
N C
CH
HC C
O O O N N
Adenine
HO P O P O P O CH2
O
O O O
C C
Triphosphate H H
H C C H
}
has three phosphate groups and so is
adenosine tri(= 3) phosphate Adenosine
The role of mitochondria ATP is produced during cellular respiration (or just simply respiration). In
in respiration is discussed in eukaryote cells, most of this process occurs in organelles known as mitochondria
chapter 3, pages 82–4. (singular = mitochondrion) which form part of the cytoplasm. Mitochondria
cannot be resolved using a light microscope but can be seen with an electron
microscope (see figure 2.15). Each mitochondrion has an outer membrane and a
highly folded inner membrane. ATP is produced by reactions that occur on the
inner folded membranes. Prokaryote cells lack mitochondria.
QUICK-CHECK
9 Of what advantage is a folded inner membrane in mitochondria?
10 What is the source of ribosomal RNA (rRNA)?
11 Some ribosomes are free in cytosol; some are attached to endoplasmic
reticulum. What is the significance of this difference?
Endoplasmic
reticulum Organelles 4 and 5: endoplasmic reticulum
and Golgi complex
We saw above that the proteins made by some cells are kept inside those cells.
Examples are contractile proteins made by muscle cells and the haemoglobins
made by red blood cells. Other cells produce proteins that are released for use
outside the cells. For example, the digestive enzyme, pepsin, is produced by cells
lining the stomach and released into the stomach cavity; the protein hormone,
insulin, is made by pancreatic cells and released into the bloodstream.
Transport of substances within cells occurs through a system of channels
known as the endoplasmic reticulum (ER). Figure 2.17 shows this system of
channels in a cell (see also figure 2.16, page 45). The channel walls are formed
by membranes.
Golgi
complex
In the Golgi complex, the proteins are packaged into secretory vesicles and may
be stored in the cytosol before they eventually fuse with the plasma membrane.
The protein is then discharged from the cell by exocytosis into the surrounding
tissue fluid. The protein may be taken up by other cells close by or may pass into
the bloodstream where it is transported to other tissues around the body.
Secretory
Rough vesicle
endoplasmic
reticulum
Membrane
fusion
occurring
Ribosomes
Transition
vesicle
Golgi
complex
QUICK-CHECK
12 Name three substances that would be produced at the surface of the
ER of a cell and transported for use outside the cell.
Lysosomes
Peroxisomes
Hydrogen peroxide (H2O2) is a product of many biochemical processes within
cells. If allowed to accumulate, it is a poisonous substance. Peroxisomes are small
membrane-bound organelles rich in the enzymes catalase and urate oxidase. The
accumulation of hydrogen peroxide is prevented by the action of catalase.
2H2O2 catalase 2H2O + O2
Separate Web
toes between
toes
Figure 2.21 In a chicken Peroxisomes detoxify various toxic materials that enter the bloodstream.
embryo, cell death brought about For example, about 25 per cent of any alcohol consumed is detoxified through
by lysosomes produces separate oxidation to aetaldehyde. Peroxisomes in different types of cells may contain
digits. Blue areas are regions where different sets of enzymes. Plant and animal cells have peroxisomes.
cell death occurs. In contrast, in
a duck embryo, cells between the Endosomes
digits do not die but are retained as
webbing. Endosomes are membrane-bound organelles found in animal cells. When material
enters a cell by endocytosis, endosomes pass on the newly ingested material to
lysosomes for digestion.
KEY IDEAS
• Lysosomes are membrane-bound sacs containing dissolved digestive
enzymes.
• Lysosomes can digest material brought into their sacs.
• Peroxisomes contain enzymes that destroy toxic materials.
• Endosomes, found in animal cells, pass on material to lysosomes for
digestion.
QUICK-CHECK
13 Lysosomes are sometimes called ‘suicide bags’. Suggest why this name
is given.
14 How is the hydrogen peroxide produced in cellular metabolism
detoxified?
15 What is the function of endosomes?
Figure 2.22
(a) Transmission
electron micrograph of
chloroplasts from the
leaf of a pea plant
(b) A three-dimensional
Outer representation of a
membrane
Stroma chloroplast
KEY IDEAS
• Chloroplasts are relatively large organelles found in photosynthetic
cells of plants and algae.
• Chloroplasts have an external membrane and layers of folded internal
membranes.
• Chlorophyll is located inside the grana of chloroplasts.
• Chloroplasts can capture the radiant energy of sunlight and convert it
to chemical energy in sugars.
QUICK-CHECK
16 What is the function of chlorophyll?
17 What are (a) thylakoids; (b) grana; (c) stroma?
Structure Function
plasma membrane Structure that controls the entry of raw materials, such as amino acids, into the cell
nucleus Organelle that has coded instructions for making the protein
ribosome Organelle where amino acids are linked, according to instructions, to build the protein
mitochondrion Organelle where ATP is formed; provides an energy source for the protein-manufacturing
activity
endoplasmic reticulum Channels through which the newly made protein is moved within the cell
Golgi complex Organelle which packages the protein into vesicles for transport across the plasma
membrane and out of the cell
(a) (b)
Protein Nucleus
Cytosol filament Nuclear Nucleolus
Endosome Cytosol
envelope
Mitochondrion
Plasma
membrane
Nucleus
Mitochondrion Ribosome
Lysosome
Nuclear Endoplasmic
envelope reticulum
Golgi Plasma
Nucleolus apparatus membrane
Ribosome
Endoplasmic Vesicle
reticulum Peroxisome Cell wall
Lysosome
Microtubule
Centriole
Peroxisome Vacuole
Protein Chloroplast
microtubule
Filament
Golgi apparatus
Vesicle
Figure 2.23 The structure and organelles of (a) an animal cell, and (b) a plant cell
Communicating
junction
Pipeline between
adjacent cells
Cell 1 Cell 2
Plasma Plasma
membrane membrane
Communicating junctions
Communicating junctions are also called gap junctions. They consist of protein-
lined pores in the membranes of adjacent cells. The proteins are aligned rather
Figure 2.26 Communicating like a series of rods in a circle with a gap down the centre (see figure 2.26)
junction of animal cells. Note the Communicating junctions permit the passage of salt ions, sugars, amino acids
pore formed by protein molecules and other small molecules as well as electrical signals from one cell to another.
aligned as if on the circumference One example of the latter is the control of the beating of the heart. A small area of
of a circle. your heart, called the pace maker, receives an electrical impulse. This electrical
impulse is transmitted to all cells of the heart through communication junctions
so that the heart ‘beats as one’.
Cytosol
Cell walls
of adjacent
plant cells
Plasmodesmata
Figure 2.28 Plasmodesmata, the junctions between plant cells. Note the relative
thickness of the section containing the walls of two plant cells, the continuation of the
cell membrane from one cell to another and the connections between smooth endoplasmic
reticulum of adjacent cells.
KEY IDEAS
• Organelles interact to facilitate the production of proteins and the
transport of these and other compounds throughout a cell.
• Cells have an internal support system called the cytoskeleton.
• In multicellular animals, some cells have connections that allow
communication with adjacent cells.
• In multicellular plants, all cells have connections that allow
communication with adjacent cells.
QUICK-CHECK
18 Name the different structures that make up the cytoskeleton of a cell.
19 List the three types of connections possible between two animal cells
and name a characteristic of each.
20 What are the connections between two plant cells called?
2
A This image shows plasmodesmata connections
between two cells. A number of cell organelles
are also visible.
B C a Is this an image of animal or plant tissue?
b Name the structures labelled A, B, C,
D and E.
c What is the function of structure F?
d What is the function of plasmodesmata?
Explain their importance.
F
E
D
3
A B This image shows a portion of a cell and some
of its organelles.
a Name the structures labelled A and B.
b Name the structure labelled C. What is its
function?
c Structures C and D are the same kind of
organelle yet their appearance is quite
different. Explain why they look so different
from each other.
C D
Questions
1 Making connections between concepts ➧ Use at least six of the key words
from this chapter to construct a concept map.
2 Analysing information and drawing conclusions ➧ Figure 2.29 is a coloured
transmission electron micrograph of a plasma cell. One function of plasma
cells is to secrete antibodies during an immune response. Note the extensive
network of endoplasmic reticulum.
a Explain whether you would expect the ER to be rough or smooth.
b Given the function of plasma cells, what other organelle would you expect
to be rather prominent in parts of this cell?
c What is the darkly stained material in the nucleus?
Figure 2.29 Transmission electron 3 Making connections between concepts ➧ Mitochondria and chloroplasts both
micrograph of a plasma cell contain circular molecules of DNA and free ribosomes. What conclusions
can reasonably be made on the basis of the presence of such structures?
4 Applying knowledge and understanding ➧ Examine table 2.2 on page 38.
a What is the difference in structure between rough and smooth endoplasmic
Fold of inner
membrane
reticulum?
b Which kind of cell shown in the table has the greater percentage of rough
endoplasmic reticulum? Which has the greater percentage of smooth
Holes in
membrane
endoplasmic reticulum?
Stalked c As a result of this difference, what would you conclude about the fate of
particle
the majority of protein produced by each cell? Explain your conclusion.
Inner membrane
5 Analysing information and drawing conclusions ➧ The folded internal
Outer membrane
membranes of mitochondria have many stalked particles on their innermost
surfaces (see figure 2.30). Given the function of mitochondria and where
Figure 2.30 Internal membrane of most of the reactions occur, of what advantage might the presence of these
mitochondria particles be for the production of ATP in the organelle?
Z
Figure 2.32
(a) Chylomicron (b) Low-density lipoprotein (LDL) (c) High-density lipoprotein (HDL)