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ICS1.
1
Genetics
1.1
Michael
Deyholos
30
November,
2010
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Figure 1.8 Chemical
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nucleotides in a fragment
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bases A,C,G,T are
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Table 1.1 Measures of genome size in selected organisms. The DNA content (1C) is shown in millions of
basepairs (Mb). Average gene density is the mean number of non-coding bases (in bp) between genes in the
genome. For eukaryotes, the chromosome number is the chromosomes counted in a gamete (1N) from each
organism.
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Chapter 2 CHROMOSOMES, MITOSIS, AND MEIOSIS
Chromosomes contain genetic information. We often take this fact for
granted, but just over a century ago, even the best biologists in the
world were uncertain of the function of these rod‐shaped structures.
We now know that most chromosomes contain a single molecule of
double‐stranded DNA that is complexed with proteins. This
arrangement allows very long DNA molecules to be compacted into a
small volume that can more easily be moved during mitosis and meiosis
(Fig 2.1). The compact structure also makes it easier for pairs of
chromosomes to align with each other during meiosis. Finally, we shall
see that chromosomal structure can affect whether genes are active or
silent.
CHROMOSOMES MAY BE LOOSE OR COMPACT
If stretched to its full length, the DNA molecule of the largest human
chromosome would be 85mm. Yet during mitosis and meiosis, this
DNA molecule is compacted into a chromosome approximately 5µm
long. Although this compaction makes it easier to transport DNA within
a dividing cell, it also makes DNA less accessible for other cellular
functions such as DNA synthesis and transcription. Thus,
chromosomes vary in how tightly DNA is packaged, depending on the
stage of the cell cycle and also depending on the level of gene activity
required in any particular region of the chromosome.
compact it. First, proteins called the core histones act as spool around
which DNA is coiled twice to form a structure called the nucleosome.
Nucleosomes are formed at regular intervals along the DNA strand,
giving the molecule the appearance of “beads on a string”. At the next
level of organization, histone H1 helps to compact the DNA strand and
its nucleosomes into a 30nm fibre. Subsequent levels of organization
involve the addition of scaffold proteins that wind the 30nm fibre into
coils, which are in turn wound around other scaffold proteins.
C h a p t e r 2 | 23
It is useful to describe the similarity between chromosomes using
Figure 2.4 appropriate terminology (Fig 2.4). Homologous chromosomes are typically
Relationships between pairs of similar, but non‐identical, chromosomes in which one member of
chromosomes and the pair comes from the male parent, and the other comes from the female
chromatids. parent. Homologs contain the same genes but not necessarily the same
alleles. Nonhomologous chromosomes contain different sets of genes, and
may or may not be distinguishable based on cytological features such as
length and centromere position. Within a chromosomes that has undergone
replication, there are sister chromatids, which are physically connected to
each other at the centromere and remain joined until cell division. Because
a pair of sister chromatids is produced by the replication of a single DNA
molecule, their sequences are essentially identical. On the other hand, non
sister chromatids come from two separate, but homologous chromosomes,
and therefore usually contain the same genes in the same order, but do not
necessarily have identical DNA sequences.
Figure 2.5
Top: FISH (Fluorescence in situ
hybridization) labeling of all 24
different human chromosomes
(1 - 22, X, and Y) in a fibroblast
nucleus, each with a different
combination of in total seven
fluorochromes.
Bottom: False color
representation of all
chromosome territories visible
in this mid-section after
computer classification.
C h a p t e r 2 | 24
MITOSIS
Cell division is essential to asexual reproduction and the development
of multicellular organisms. Accordingly, the primary function of mitosis
is to ensure that each daughter cell inherits identical genetic material,
i.e. exactly one copy of each chromosome. To make this happen,
replicated chromosomes condense (prophase), and are positioned
near the middle of the dividing cell (metaphase), and then one sister
chromatid from each chromosome migrates towards opposite poles of
the dividing cell (anaphase), until the identical sets of chromosomes
are completely separated from each other within the newly formed
nuclei of each daughter cell (telophase) (see Figs. 2.5‐2.7 for diagrams
of the process). This is followed by the completion of the division of
the cytoplasm (cytokinesis). The movement of chromosomes is aided
by microtubules that attach to the chromosomes at centromere.
MEIOSIS
Meiosis, like mitosis, is also a necessary part of cell division. However,
in meiosis not only do sister chromatids separate from each other,
homologous chromosomes also separate from each other. This extra,
reductional step of meiosis is essential to sexual reproduction. Without
meiosis, the chromosome number would double in each generation of a
species and would quickly become too large to be viable.
Meiosis is divided into two stages designated by the roman numerals I
and II. Meiosis I is called a reductional division, because it reduces the
number of chromosomes inherited by each of the daughter cells.
Meiosis I is further divided into Prophase I, Metaphase I, Anaphase I,
and Telophase I, which are roughly similar to the corresponding stages
of mitosis, except that in Prophase I and Metaphase I, homologous
chromosomes pair with each other in transient structures called
bivalents (Figs. 2.7, 2.8). This is an important difference between
mitosis and meiosis, because it affects the segregation of alleles, and
also allows for recombination to occur through crossing‐over, as
described later in the course. During Anaphase I, one member of each
pair of homologous chromosomes migrates into a daughter cell.
Meiosis II is essentially the same as mitosis, with one sister chromatid
from each chromosome separating to produce two identical cells.
Because Meiosis II, like mitosis, results in products that contain
identical sequences, Meiosis II is called an equational division.
C h a p t e r 2 | 25
Figure 2.7 Mitosis and meiosis. Note the similarities and differences between metaphase in
mitosis and metaphase I and II of meiosis.
Figure 2.8 Meiosis in Arabidopsis (n=5). Panels A-C show different stages of prophase I, each with an
increasing degree of chromosome condensation. Subsequent phases are shown: metaphase I (D), telophase I
(E), metaphase II (F), anaphase II (G), and telophase II (H).
C h a p t e r 2 | 26
THE CELL CYCLE AND CHANGES IN DNA CONTENT
The life cycle of an eukaryotic cell can generally be divided into at least
four stages (Fig. 2.9). When a cell is produced through fertilization or
cell division, there is usually a lag before it undergoes DNA synthesis.
This lag period is called Gap 1 (G1), and ends with the onset of the DNA
synthesis (S) phase, during which each chromosome is replicated.
Following replication, there may be another lag, called Gap 2 (G2),
Figure 2.9 A typical before mitosis (M). Cells undergoing meiosis do not usually have a G2
eukaryotic cell cycle.
phase. Interphase is as term used to describe all phases of the cell
cycle excluding mitosis or meiosis. A typical cell cycle is shown in Fig.
2.9. My variants of this generalized cell cycle also exist. Some cells
never leave G1 phase, and are said to enter a permanent, non‐dividing
stage called G0. On the other hand, some cells undergo many rounds of
DNA synthesis (S) without any mitosis or cell division. These
endoreduplicated cells are described later in this chapter.
Understanding the control of the cell cycle is an active area of research,
particularly because of the relationship between cell division and
cancer.
The amount of DNA within a cell changes following each of the
following events: fertilization, DNA synthesis, mitosis, and meiosis (Fig
2.10). We use “c” to represent the DNA content in a cell, and “n” to
represent the number of complete sets of chromosomes. In a gamete
(i.e. sperm or egg), the amount of DNA is 1c, and the number of
chromosomes is 1n. Upon fertilization, both the DNA content and the
number of chromosomes doubles to 2c and 2n, respectively. Following
DNA synthesis, the DNA content doubles again to 4c, but each pair of
sister chromatids is still counted as a single chromosome, so the
number of chromosomes remains unchanged at 2n. If the cell
undergoes mitosis, each daughter cell will be 2c and 2n, because it will
receive half of the DNA, and one of each pair of sister chromatids. In
C h a p t e r 2 | 27
contrast, the cells that are produced from the meiosis of a 2n, 4c cell are
1c and 1n, since each pair of sister chromatids, and each pair of
homologous chromosomes divides during meiosis.
Figure 2.10
Changes in DNA
and chromosome
content during
the cell cycle.
For simplicity,
nuclear
membranes are
not shown, and
all chromosomes
are represented
in a similar stage
of condensation.
C h a p t e r 2 | 28
KARYOTYPES SHOW CHROMOSOME NUMBER AND STRUCTURE
Each eukaryotic species has its total nuclear genome divided among a
number of chromosomes that is characteristic of that species. For
example, a haploid human nucleus (i.e. sperm or egg) normally has 23
chromosomes (n=23), and a diploid human nucleus has 23 pairs of
chromosomes (2n=46). Various stains and fluorescent dyes produce
characteristic banding patterns on some chromosomes. This can make
it easier to identify specific chromosomes. The number of
chromosomes varies between species (see Table 1.1), but there appears
to be very little correlation between chromosome number and either
the complexity of an organism or its total amount genomic DNA.
The most familiar human aneuploidy is trisomy‐21 (i.e. three copies of
chromosome 21), which is one cause of Down’s syndrome. Most (but
not all) other human aneuploidies are lethal at an early stage of
C h a p t e r 2 | 29
embryonic development. Note that aneuploidy usually affects only one
set of homologs within a karyotype, and is therefore distinct from
polyploidy, in which the entire karyotype is duplicated (see below).
Aneuploidy is almost always deleterious, whereas polyploidy appears
to be beneficial in some organisms, particularly some species of plants.
Figure 2.12
Structural abberations
in chromosomes.
Structural defects in chromosomes are another type of abnormality that
can be detected in karyotypes (Fig 2.12). These defects include
deletions, duplications, and inversions, which all involve changes in a
segment of a single chromosome. Insertions and translocations
involve two non‐homologous chromosomes. In an insertion, DNA from
one chromosome is unidirectional, while in translocation, the transfer
of chromosomal segments is bidirectional. Structural defects affect
only part of a chromosome, and so tend to be less harmful than
aneuploidy. In fact, there are many examples of ancient chromosomal
rearrangements in the genomes of species including our own.
Duplications of some small chromosomal segments, in particular, may
have some evolutionary advantage by providing extra copies of some
genes, which can then evolve in new ways.
C h a p t e r 2 | 210
POLYPLOIDY
Humans, like most animals and all of the eukaryotic genetic model
organisms in wide use, are diploid. This means that most of their cells
have two homologous copies of each chromosome. In contrast, many
plant species and even a few animal species are polyploids. This
means there are more than two homologs of each chromosome in each
cell.
When describing polyploids, we use the letter “x” to define the level of
ploidy. A diploid is 2x, because there are two basic sets of
chromosomes, and a tetraploid is 4x, because it contains four copies of
each chromosome. For clarity when discussing polyploids, geneticists
will often combine the “x” notation with the “n” notation already
defined previously in this chapter. Thus for both diploids and
polyploids, “n” is the number of chromosomes in a gamete, and “2n” is
the number of chromosomes following fertilization. For a diploid,
therefore, n=x, and 2n=2x. For a tetraploid, n=2x, and 2n=4x.
Like diploids (2n=2x), stable polyploids generally have an even number
of copies of each chromosome: tetraploid (2n=4x), hexaploid (2n=6x),
and so on. The reason for this is clear from a consideration of meiosis.
Remembering that the purpose of meiosis is to reduce the sum of the
genetic material by half, meiosis can equally divide an even number of
chromosome sets, but not an odd number. Thus, polyploids with an
uneven number of chromosomes (e.g. triploids, 2n=3x) tend to be
sterile, even if they are otherwise healthy. The mechanism of meiosis in
stable polyploids is essentially the same as in diploids: during
metaphase I, homologous chromosomes pair with each other.
Depending on the species, all of the homologs may be aligned together
at metaphase, or in multiple separate pairs. For example, in a
tetraploid, some species may form tetravalents in which the four
homologs from each chromosome align together, or alternatively, two
pairs of homologs may form two bivalents. Note that because that
mitosis does not involve any pairing of homologous chromosomes,
mitosis is equally effective in diploids, even‐number polyploids, and
odd‐number polyploids.
Triploidy is used in the production of seedless fruits, such as
watermelon, grapes and bananas. All of the tissues of these fruit are
triploid. Because almost all of the cells of the plant, including its fruit,
are produced through mitosis, the uneven number of chromosome sets
does not affect their development. However, cells that contribute to the
production of gametes are produced through meiosis, and because the
triploids are unable to complete normal meioses, their gametes fail to
develop, so no zygotes are formed, and the seeds (which normally
contain embryos that develop from the zygotes) are aborted.
C h a p t e r 2 | 211
Figure 2.14.
Endoreduplicated
chromosomes from an
insect salivary gland.
The banding pattern is
produced with
fluorescent labels.
ENDOREDUPLICATION
Endoreduplication, also known as endopolyploidy, is a special type
of tissue‐specific genome amplification that occurs in many types of
plant cells and in specialized cells of some animals including humans.
Endoreduplication does not affect the germline or gametes, so species
with endopolyploidy are not considered polyploids. Endopolyploidy
C h a p t e r 2 | 212
occurs when a cell undergoes multiple rounds of DNA synthesis (S‐
phase) without any mitosis. This produces multiple chromatids of each
chromosome. Endopolyploidy seems to be associated with cells that
are metabolically very active, and produce a lot of enzymes and other
proteins in a short amount of time. The highly endoreduplicated
salivary gland chromosomes of D. melanogaster have been useful
research models in genetics, since their relatively large size makes
them easy to study under the microscope.
GENE BALANCE
Why do trisomies, duplications, and other chromosomal abnormalities
that increase gene copy number sometimes have a negative effect on
the normal development or physiology of an organism? This is
particularly intriguing because in many species, aneuploidy is
detrimental or lethal, while polyploidy is tolerated or even beneficial.
The answer is probably related to the concept of gene balance, which
can be summarized as follows: genes, and the proteins they produce,
have evolved to be part of complex metabolic and regulatory networks.
Some of these networks function best when certain enzymes and
regulators are present in specific ratios to each other. Increasing the
gene copy number for just one part of the network may throw the
network out of balance, leading to increases or decreases of certain
metabolites, which may be toxic in high concentrations or which may
be limiting in other important processes in the cell. The activity of
genes and metabolic networks is regulated in many different ways
besides changes in gene copy number, so duplication of just a few genes
will usually not be harmful. However, trisomy and large segmental
duplications of chromosomes affect the dosage of so many genes that
cellular networks are unable to adjust to the changes.
ORGANELLAR GENOMES
Chromosomes also exist outside of the nucleus, within both the
chloroplast and mitochondria. These organelles are likely the remnants
of a prokaryotic endosymbionts that entered the cytoplasm of ancient
progenitors of today’s eukaryotes. These endosymbionts had their
own, circular chromosomes, like most bacteria that exist today.
Likewise, chloroplasts and mitochondria also have circular
chromosomes that behave more like bacterial chromosomes than
eukaryotic chromosomes, i.e. these organellar genomes do not undergo
mitosis or meiosis. Organellar genomes are also often present in
multiple copies within each organelle, and in most species are inherited
maternally.
C h a p t e r 2 | 213
_______________________________________________________________________________________
SUMMARY
• Chromosomes are complex and dynamic structures consisting of DNA and proteins
(chromatin).
• The degree of chromatin compaction varies between heterochromatic and euchromatic
regions and between stages of the cell cycle.
• Some chromosomes can be distinguished cytologicaly based on their length, centromere
position, and banding patterns when stained dyes or labelled with sequence‐specific
probes.
• Homologous chromosomes contain the same genes, but not necessarily the same alleles.
Sister chromatids usually contain the same genes and the same alleles.
• Mitosis reduces the c‐number, but not the n‐number. Meiosis reduces both c and n.
• Homologous chromosomes associate with each other during meiosis, but not mitosis.
• Several types of structural defects in chromosomes occur naturally, and can affect cellular
function and even evolution.
• Aneuploidy results from the addition or subtraction of one or more chromosomes from a
group of homologs, and is usually deleterious to the cell.
• Polyploidy is the presence of more than two complete sets of chromosomes in a genome.
Even‐numbered multiple sets of chromosomes can be stably inherited in some species,
especially plants.
• Endopolyploidy is tissue‐specific type of polyploidy observed in some species, including
diploids.
• Both aneuploidy and structural defects such as duplications can affect gene balance.
• Organelles also contain chromosomes, but these are much more like prokaryotic
chromosomes than the nuclear chromosomes of eukaryotes.
C h a p t e r 2 | 214
KEY TERMS
chromosome mitosis c
core histones prophase karyotype
nucleosome metaphase aneuploidy
30nm fiber anaphase monsomic
histone H1 telophase trisomic
scaffold proteins prophase Down’s syndrome
heterochromatin meiosis deletion
euchromatin prophase (I, II) duplication
microsatellite metaphase (I, II) insertion
chromatid anaphase (I, II) inversion
centromere telophase (I, II) translocation
metacentric cytokinesis non‐disjunction
acrocentric bivalent chromosome breakage
telocentric reductional division polyploid
holocentric equational division x
telomere G 1 tetravalent
homolog G 2 endoreduplication
non‐homologous S endopolyploidy
chromatid M gene balance
sister chromatid G 0 cellular network
non‐sister chromatid interphase endosymbiont
interphase n organellarchromo
_______________________________________________________________________________________
STUDY QUESTIONS
2.1 Define chromatin. What is the difference 2.4
between DNA, chromatin and a) How many centromeres does a typical
chromosomes? chromosome have?
b) What would happen if there was more
2.2 Species A has n=4 chromosomes and than one centromere per chromosome?
Species B has n=6 chromosomes. Can you c) What if a chromosome had zero
tell from this information which species has centromeres?
more DNA? Can you tell which
species has more genes? 2.5 For a diploid with 2n=16 chromosomes,
how many chromosomes and chromatids
2.3 The answer to question 2 implies that are per cell present in the gamete, and
not all DNA within a chromosome encodes zygote and immediately following G1, S, G2,
genes. Can you name any examples of mitosis, and meiosis?
chromosomal regions that contain
relatively few genes? 2.6 Bread wheat (Triticum aestivum) is a
hexaploid. Using the nomenclature
presented in class, an egg cell of wheat has
!
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C h a p t e r 4 | 45
Box 4‐1 Transposable Elements
Transposable elements (TEs) occur naturally throughout the chromosomes of almost all
organisms. These DNA sequences have a unique ability to be inserted into new locations in the
genome, after being cut or copied from their original location. TEs are also known as mobile
genetic elements, or more informally as jumping genes. The locations into which TEs are
inserted are not entirely random, but TEs can in principle be inserted into almost any region of
the genome. TEs can therefore move into other genes, causing mutations. Researchers have
methods of artificially increasing the rate of transposition, making TEs a useful type of mutagen.
However the biological importance of TEs extends far beyond their use in mutant screening; TEs
are also important causes of disease and phenotypic instability, and they are a major force in
evolution.
There are two major classes of TEs in eukaryotes (Figure 4‐B1). Class I elements include
retroposons and retrotransposons; these are copied by means of an RNA intermediate. The
transcript is reverse transcribed into DNA before being inserted elsewhere in the genome
through the action of enzymes such as integrase. Class II elements are known also as
transposons; these do not use reverse transcriptase or an RNA intermediate for transposition.
Using an enzyme called transposase, most transposons are cut from their original location and
then this excised dsDNA fragment is inserted into a new location. Note that the name transposon
is sometimes used incorrectly to refer to any type of TEs, but in this book we use transposon to
refer only to Class II elements.
Figure 4B1 . Representative examples of the two main types of transposable elements. (TEs)
Class I elements transpose via an ssRNA intermediate, which is reverse transcribed to dsDNA
prior to insertion of this copy in a new site in the genome. Class II elements do not involve an
RNA intermediate; most Class II elements are cut from their original location as dsDNA, prior to
being inserted into a new site in the genome. Although the diagram shows TEs being inserted on
the same chromosome as they originated from, TEs can also move to other chromosomes within
the same cell.
C h a p t e r 4 | 46
TEs are relatively short DNA sequences (between 100bp and 10kb), and encode no more than a
few proteins (if any). Normally, the protein‐coding genes within a TE are all related to the TE’s
own transposition functions, e.g. reverse transcriptase, transposase, and integrase. However,
some TEs (of either Class I or II) do not encode any proteins at all. These nonautonomous TEs
can only transpose if they are supplied with enzymes produced by other, autonomous TEs
located elsewhere in the genome. In all cases, enzymes for transposition recognize conserved
nucleotide sequences within the TE, which show the enzymes where to begin cutting or copying,
and re‐insertion.
The human genome is nearly 45% TEs, the vast majority of which are families of Class I elements
called LINEs and SINEs. The short, Alu type of SINE occurs in more than one million copies in
the human genome (compare this to the approximately 30,000, non‐TE, protein‐coding genes in
humans). Indeed, TEs make up a significant portion of the genomes of almost all eukaryotes.
Class I elements, which usually transpose via a copy‐and‐paste mechanism, tend to be more
abundant than Class II elements, which mostly use a cut‐and‐paste mechanism. But even the
cut‐paste mechanism can lead to an increase in TE copy number, in some circumstances (for
example, if the site vacated by the excised transposon is repaired with a DNA template from a
homologous chromosome that itself contains a copy of a transposon).
Besides greatly expanding the DNA content of genomes, TEs contribute to genome evolution in
many other ways. As already mentioned, they may disrupt gene function by insertion into a
gene’s coding region or regulatory region. More interestingly adjacent regions of chromosomal
DNA are sometimes mistakenly transposed along with the TE; this can lead to gene duplication.
The duplicated genes are then free to evolve independently, leading in some cases to the
development of new functions. The breakage of strands by TE excision and integration can
disrupt genes, and can lead to chromosome rearrangement or deletion if errors are made during
strand rejoining. Furthermore, having so many similar TE sequences distributed throughout a
chromosome sometimes allows mispairing of regions of homologous chromosomes at meiosis,
which can cause unequal crossing‐over, resulting in deletion or duplication of large segments of
chromosomes. Thus, TEs are an important evolutionary force, and are not merely “junk DNA” as
they were once called.
Figure 4B2. Barbara McClintock won a Nobel Prize for her discovery of TEs. She did so by
studying pigment variegation in maize kernels, which is caused by the movement of TEs in and
out of pigmentation genes. This is an example of phenotypic instability.
C h a p t e r 4 | 47
The new strand therefore bears a mutation, which will be inherited, in
all the strands that are subsequently replicated from it.
Intercalating agents are another type of
chemical mutagen. These induce mutations by
inserting between the stacked bases at the
center of the DNA helix (Figure 4.7). This
intercalation distorts the shape of the DNA helix,
which can cause the wrong bases to be added to
a growing DNA strand during DNA synthesis.
Intercalating agents tend to be flat, planar
molecules such as benzopyrene, a component
of wood and tobacco smoke. Another important
intercalating agent is thalidomide, an anti‐
nausea drug whose harmful effects were
unknown until its consumption by thousands of
Figure 4.7 Benzopyrene
pregnant woman resulted in birth defects. (circled in red) is an
Finally ethidium bromide, the dye that example of an
fluorescently stains DNA in laboratory assays, is intercalating agent
also an intercalating agent. For this reason,
molecular biologists are trained to handle this chemical carefully.
PHYSICAL
Anything that damages DNA by transferring energy to it can be
considered a physical mutagen. Usually this involves radioactive
particles, x‐rays, or UV light. Smaller, fast moving particles may
substitute or delete a single base, while larger, slightly slower particles
induce larger deletions by breaking the double stranded helix. Physical
C h a p t e r 4 | 48
mutagens can also create unusual structures in DNA, such as the
thymine dimers formed by UV light (Figure 4.8). Thymine dimers
disrupt normal base‐pairing in the double helix, and may block
replication altogether if not repaired by the cell.
MUTANT SCREENING: FORWARD GENETICS
One way to identify genes that affect a particular biological process is to
induce random mutations in a population, and then look for individuals
with phenotypes that might be caused by a disruption of a particular
biochemical pathway. This strategy of mutant screening has been
used very effectively to better understand the molecular components of
hundreds of different biological processes. For example, to better
understand processes of memory and learning, researchers have
screened mutagenized populations of Drosophila to identify flies (or
larvae) that lack the normal ability to learn to associate a particular
odor with an electric shock. Some of the genes identified by this mutant
screen may be relevant to learning and memory in other animals,
including conditions such as Alzheimer’s disease in humans.
Exposure of an organism to a mutagen causes mutations in essentially
random positions along the chromosomes. Most of the mutant
phenotypes recovered from a genetic screen are caused by lossof
function mutations. These are changes in the sequence of an allele that
cause it to no longer produce the same level of active protein as the
wild‐type allele. Loss‐of‐function alleles tend to be recessive because
the wild‐type allele is haplosufficient (see Chapter 6). A loss‐of‐
function allele that produces no active protein is called an amorph, or
null. On the other hand, alleles with only a partial loss‐of‐function are
called hypomorphic. More rarely, a mutant allele may have a gainof
function, producing either more of the active protein (hypermorph)
or producing an active protein with a new function (neomorph).
C h a p t e r 4 | 49
In a typical mutant screen, researchers will treat a parental population
with a mutagen. This may, for example, involve soaking seeds in EMS,
or mixing this mutagen with the food fed to flies. The individuals that
are directly exposed with the mutagen are called the M0 generation. No
phenotypes will be visible among the M0 generation, in part because not
all somatic (i.e. non‐reproductive) cells of the M1 individuals will be
affected in the same way. More important in the M1 generation are the
germline cells: gametes and any of their developmental precursors.
Since a single gamete contributes half of the chromosomes to every cell
of the next (M2) generation, mutagenizing gametes or the cells that
produce them is the easiest way to generate individuals in which every
cell bears the same mutation. In most cases, however, the M1
individual will be heterozygous for any induced mutation; this is
because it would be very unlikely for the same gene to have been
mutated in the other gamete that contributed to the M1 individual.
Because most induced mutations are recessive, the M1 generation must
therefore be selfed (i.e. crossed to siblings, or self‐fertilization if the
species is a hermaphrodite like worms and most plants), and the
following generations (e.g. M2, in which mutant alleles could potentially
become homozygous) examined for the presence of novel phenotypes.
Once a relevant mutant has been identified, geneticists can begin to
make inferences about what the normal function of the mutated gene is,
based on its mutant phenotype.
SOME MUTATIONS MAY NOT HAVE DETECTABLE PHENOTYPES
The vast majority of mutations (especially substitutions) have no effect
on the phenotype. Often, this is because the mutation is silent; it
changes the DNA sequence of a non‐coding region of the DNA, or else
the changes a base within a codon without changing the amino acid that
it encodes (recall that the genetic code is degenerate; for example, GCT,
GCC, GCA, and GCG all encode alanine). There are also cases where a
mutation can cause a complete loss‐of‐function of a gene, yet not
produce a phenotype even when the mutant allele is homozygous. This
can often be attributed to genetic redundancy, i.e. the encoding of
similar genes at more than one locus in the genome. It is important
therefore to remember this important limitation of mutational analysis:
genes with redundant functions cannot be easily identified by mutant
screening.
C h a p t e r 4 | 410
phenotypic class, giving a phenotypic ratio of 1:0 (which is the same as
3:0).
Many genes are first identified in mutant screens, and so they tend to
be named after their mutant phenotypes. This can cause some
confusion for students of genetics. For example, we have already
encountered an X‐linked gene named white in fruit flies. Null mutants
of white have white eyes, but the normal function of the white gene is
actually the production of a red pigment.
COMPLEMENTATION TESTING
Mutations in different genes can produce the same phenotype. For
example, in the biochemical pathway shown in Figure 4.9, a plant that
lacks the function of gene A (genotype aa) would produce mutant,
white flowers that looked just like the flowers of a plant that lacked the
function of gene B (genotype bb). The genetics of two loci are discussed
more in the following chapters.
As explained earlier in this chapter, mutant screening is one of the main
activities of geneticists. When geneticists find two mutants with similar
phenotypes, either in natural populations or during a mutant screen, an
immediate question is whether or not the mutants have lost the
function of the same gene. The other possibility is that each mutant has
C h a p t e r 4 | 411
lost the function of a different gene. If they are both mutants of the
same gene, then their genotypes can both be represented as aa. If each
has a mutation in a different gene, then the genotype of one individual
can be represented as aa, and the other individual as bb (or more
completely as aaBB and AAbb, respectively).
A technique called complementation testing can be used to determine
whether two individuals with the same phenotype carry mutations in
the same gene or different genes. The only requirement of this test is
two pure‐breeding individuals with the same phenotype; no prior
knowledge of the genes or biochemical pathways is required. To
perform a complementation test, two homozygous individuals with
similar mutant phenotypes are crossed (Figure 4.10). If the F1 progeny
all have the mutant phenotype, then we infer that same gene is mutated
in each parents. If the F1 progeny all appear to be wild‐type, then each
of the parents most likely carries a mutation in a different gene (Figure
4.11).
Figure 4.10
In a typical
complementation test,
the genotypes of two
parents are unknown
(although they must be
pure breeding,
homozygous mutants).
If the F1 progeny all
have a mutant
phenotype there (Case
1), there is no
complementation. If
the F1 progeny are all
wild-type, the
mutations are said to
have complemented
each other. See Figure
4.11 for interpretation.
C h a p t e r 4 | 412
Figure 4.11
The pure breeding,
homozygous mutants
parents had unknown
genotypes before the
complementation test, but
it could be assumed that
they were either mutations
in the same genes (Case 1)
or different genes (Case
2). In Case 1, all of the
progeny would have a
mutant phenotype,
because they would all
have the same,
homozygous genotype as
the parents. In Case 2,
each parent has a mutation
in a different gene,
therefore none of the F1
progeny will be
homozygous mutant at
any one locus. Note that
the genotype in Case 1
could be written as either
aa or aaBB.
Thus, if two homozygous mutants produce F1 progeny that have a wild‐
type phenotype, complementation is said to have occurred, because
the wild‐type alleles of each of the genes were able to compensate for
the recessive, mutant alleles that were also inherited (Case 2, Figure
4.11). On the other hand, if the F1 progeny all have a mutant
phenotype, (case 1, Figure 4.11), the mutant genotypes fail to
complement each other, because they contain mutations of the same
gene. These could be either the same mutant alleles, or different mutant
alleles of the same gene. If the two mutants are independent (e.g. they
came from different natural populations or from independently
mutagenized individuals), the mutations are probably different alleles
of the same gene. All mutants that fail to complement each other are
said to be in the same complementation group.
C h a p t e r 4 | 413
_______________________________________________________________________________________________________________
SUMMARY
• When a variation in DNA sequence originated recently, and is rare in a population, we call
that change a mutation.
• When variations in DNA sequence co‐exist in a population, and neither one can be
meaningfully defined as wild‐type, we call the variations polymorphisms.
• Mutations may either occur spontaneously, or may be induced by exposure to mutagens.
• Mutations may result in either substitutions, deletions, or insertions.
• Mutation usually causes either a partial or complete loss of function, but sometimes
results in a gain of function, including new functions.
• Spontaneous mutations arise from many sources including natural errors in DNA
replication, usually associated with base mispairing, or else insertion deletion especially
within repetitive sequences.
• Induced mutations result from mispairing, DNA damage, or sequence interruptions
caused by chemical, biological, or physical mutagens.
• By randomly inducing mutations, then screening for a specific phenotype, it is possible to
identify genes associated with specific biological pathways.
• Transposable elements are dynamic, abundant components of eukaryotic genomes and
important forces in evolution.
• Transposable elements are dynamic, abundant components of eukaryotic genomes and
important forces in evolution.
• Mutation of different genes can produce a similar phenotype.
• Complementation testing determines whether two mutants are the result of mutation of
the same gene, or if each mutant is caused by mutation of a different gene.
• The efficiency of mutant screening is limited by silent mutations, redundancy, and
embyronic lethality.
C h a p t e r 4 | 414
KEY TERMS
mutation transposon
mutant retrotransposon loss‐of‐function
polymorphism reverse transcriptase gain‐of‐function
insertion transposase amorph
deletion non‐autonomous null
substitution autonomous hypomorph
mutagen SINE, LINE, Alu hypermorph
biological mutagen P‐element neopmorph
chemical mutagen T‐DNA somatic
physical mutagen alkylation agent germline
tautomer EMS M0, M1, M2
mispairing intercalating agent silent mutation
loop benzopyrene redundancy
SSR ethidium bromide lethality
insertional mutagen thymine dimer complementation group
Class I, Class II mutant screen
_______________________________________________________________________________________________________________
STUDY QUESTIONS
4.1 How are polymorphisms and mutations However there are twice as many females as
alike? How are they different? males in the F1 progeny of this cross.
a) Propose a hypothesis to explain these
4.2 What are all of the ways a substitution observations.
can occur in a DNA sequence? b) How could you test your hypothesis?
4.3 What are all of the ways a deletion can 4.7 You decide to use genetics to investigate
occur in a DNA sequence? how your favourite plant makes its
flowers smell good.
4.4 What are all of the ways an insertion can a) What steps will you take to identify some
occur in a DNA sequence? genes that are required for production of the
sweet floral scent? Assume that this plant
4.5 In the context of this chapter, explain the is a self‐pollinating diploid.
health hazards of smoking tobacco. b) One of the recessive mutants you
identified has fishy‐smelling flowers, so you
4.6 You have exposed a female fruit fly to a name the mutant (and the mutated gene)
mutagen. Mating this fly with a non‐ fishy. What do you hypothesize about the
mutagenized male produces offspring that normal function of the wild‐type fishy gene?
appear to be completely normal.
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blindness are inherited as XR-traits.
!! Colour blindness is diagnosed using tests
such as this Ishihara Test.
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Photo and Illustration Credits
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