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GENETICS 1.1
Genetics 1.1 Michael Deyholos 30 November, 2010 The first edition of this book was produced in January, 2009 as instructional material for students in Biology 207 at the University of Alberta, and is released to the public for non-‐commercial use under the Creative Commons License (See below). Users are encouraged to make modifications and improvements to the book. All text in the original edition was written by Michael Deyholos, Ph.D. Photos and some diagrams were obtained from various, non-‐copyrighted sources, including Flickr, Wikipedia, Public Library of Science, and Wikimedia Commons. Photo attributions are listed at the end of the book.
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blending inheritance
Mendel
not
particulateinheritance
alleles
genes
Figure 1.1 Gregor Mendel
1 2
Streptococcus pneumoniae
Griffith
Avery,MacLeod and McCarty
Figure 1.2 Inheritance of flower color in peas. Mendel observed that a cross between pure breeding,white and purple peas (generation P) produced only progeny (generation F1) with purple flowers. However, white flowered plant reappeared among the F2 generation progeny of a mating between two F1 plants. The symbols P, F1 and F2 are abbreviations for parental, first filial, and second filial generations, respectively.
Figure 1.3 Colonies of Rough (top) and Smooth (bottom) strains of S. pneumoniae.
1 3
Hershey and Chase
Escherichia coli
E. coli
Figure 1.4 Experiments of Griffith and of Avery , MacLeod and McCarty. R strains of S. pneumoniae do not cause lethality. However, DNA-containing extracts from pathogenic S strains are sufficient to make R strains pathogenic.
Figure 1.5 Electronmicrograph of T2 bacteriophage on surface of E. coli
1 4
Watson and Crick
Chargaff�’s Rules
Figure 1.6 When 32P-labeled phage infects E. coli, radioactivity is found only in the bacteria after the phage are removed by agitation and centrifugation. In contrast, after infection with 35S-labeled phage, radioactivity is found only in the supernatant that remains after the bacteria are removed.
Figure 1.7 DNA structure
1 5
Central Dogma
Figure 1.8 Chemical structure of two pairs of nucleotides in a fragment of double-stranded DNA. Sugar, phosphate, and bases A,C,G,T are labeled. Hydrogen bonds between bases on opposite strands are shown by dashed lines. Note that the G-C pair has more hydrogen bonds than A-T. The numbering of carbons within sugars is indicated by red numbers. Based on this numbering the polarity of each strand is indicated by the labels 5�’ and 3�’.
Figure 1.9 Central Dogma of molecular biology
1 6
c value
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.
Saccharomyces cerevisiae
Caenorhabditis elegans
1 7
Drosophila melanogaster
Mus musculus
Daniorerio
Arabidopsis thaliana
D. melanogaster
Figure 1.10 Some of the most important genetic model organisms in use today. Clockwise from top left: yeast, fruit fly, arabidopsis, mouse, roundworm, zebrafish.
1 8
Saccharomyces cerevisiaeCaenorhabditis elegansDrosophila melanogasterMus musculusDanio rerioArabidopsis thalianaEscherichia coli
1 9
1.1
1.2
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b)
c)
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1.3
1.4
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c)
1.5S. pneumoniae
1.6
1.7 a)
b)
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1.8a)
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1.9 a)
b)
1.10a)
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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.
There are several different levels of structural organization in eukaryotic chromosomes, with each successive level contributing to the further compaction of DNA (Fig. 2.2). For more loosely compacted DNA, only the first few levels of organization may apply. Each level involves a specific set of proteins that associate with the DNA to
Chapter 2 CHROMOSOMES, MITOSIS, AND MEIOSIS
Figure 2.1 Moving chromosomes (blue) towards the poles at anaphase requires many proteins (red), all of which interact with microtubules (green).
C h a p t e r 2 | 22
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.
Chromosomes stain very intensely with some types of dyes, which is how they got their name (chromosome means “colored body”). Certain dyes stain some regions within a chromosome more intensely that others, giving some chromosomes a banded appearance. The material that makes up chromosomes, which we now know to be proteins and DNA, is called chromatin. There are two general types of chromatin. Euchromatin is more loosely packed, and tends to contain more genes that are being transcribed, as compared to the more densely compacted heterochromatin which is rich in short, repetitive sequences called microsatellites.
Chromosomes also contain other distinctive features such as centromeres and telomeres. Both of these are heterochromatic. In most cases, each chromosome contains one centromere. These sequences are bound by centromeric proteins that link the centromere to microtubules that transport chromosomes during cell division. Under the microscope, centromeres can sometimes appear as constrictions in the body of the chromosome (Fig. 2.3). If a centromere is located near the middle of a chromosome, it is said to be metacentric, while an acrocentric centromere is closer to one end of a chromosome, and a telocentric chromsome is at the very end. More rarely, in a holocentric centromere, no single centromere can be defined and the entire chromsome acts as the centromere. Telomeres are repetitive sequences near the ends of linear chromosomes, and are important in maintaining the length of the chromosomes during replication, and protecting the ends of the chromosomes from alterations.
Figure 2.2 Successive stages of chromosome compaction depend on the introduction of additional proteins.
Figure 2.3 A pair of metacentric chromosomes. The arrow shows a centromeric region.
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It is useful to describe the similarity between chromosomes using appropriate terminology (Fig 2.4). Homologous chromosomes are typically pairs of similar, but non‐identical, chromosomes in which one member of the pair comes from the male parent, and the other comes from the female 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, nonsister 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 2.4 Relationships between chromosomes and chromatids.
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.
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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.
Figure 2.6 Mitosis in arabidopsis showing fluorescently labeled chromosomes (blue) and microtubules (green) at metaphase, anaphase and telophase (from left to right).
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), before mitosis (M). Cells undergoing meiosis do not usually have a G2 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
Figure 2.9 A typical eukaryotic cell cycle.
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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.
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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.
A karyotype shows the complete set of chromosomes of an individual (Fig. 2.11). Analysis of karyotypes can identify of chromosomal abnormalities, including aneuploidy, which is the addition or subtraction of a chromosome from a pair of homologs. More specifically, the absence of one member of a pair of homologous chromosomes is called monosomy. On the other hand, in a trisomy, there are three, rather than two homologs of a particular chromosome. Different types of aneuploidy are sometimes represented symbolically; if 2n symbolizes the normal number of chromosomes in a cell, then 2n1 indicates monosomy and 2n+1 represents trisomy.
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
Figure 2.11 Karyotype of a normal human male
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.
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.
Chromosomal abnormalities arise in many different ways. Many of these can be traced to rare errors in natural cellular processes. Nondisjunction is the failure of at least one pair of chromosomes or chromatids to separate during mitosis or meiosis. Chromosome breakage also occurs infrequently as the result of physical damage (such as radiation), movement of some types of transposons, and other factors. During the repair of a broken chromosome, deletions, insertions, translocations and even inversions can be introduced.
Figure 2.12 Structural abberations in chromosomes.
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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.
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If triploids cannot make seeds, how do we obtain enough triploid individuals for cultivation? The answer depends on the plant species involved. In some cases, such as banana, it is possible to propagate the plant asexually; new progeny can simply be grown from cuttings from a triploid plant. On the other hand, seeds for seedless watermelon are produced sexually: a tetraploid watermelon plant is crossed with a diploid watermelon plant. Both the tetraploid and the diploid are fully fertile, and produce gametes with two (1n=2x) or one (1n=1x) sets of chromosomes, respectively. These gametes fuse to produce a zygote (2n=3x), that is able to develop normally into an adult plant through multiple rounds of mitosis, but is unable to compete normal meiosis or produce seeds.
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
Figure 2.13. Part of a triploid watermelon, showing white, aborted seeds within the flesh
Figure 2.14. Endoreduplicated chromosomes from an insect salivary gland. The banding pattern is produced with fluorescent labels.
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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.
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_______________________________________________________________________________________
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.
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KEY TERMS
chromosome core histones nucleosome 30nm fiber histone H1 scaffold proteins heterochromatin euchromatin microsatellite chromatid centromere metacentric acrocentric telocentric holocentric telomere homolog non‐homologous chromatid sister chromatid non‐sister chromatid interphase
mitosis prophase metaphase anaphase telophase prophase meiosis prophase (I, II) metaphase (I, II) anaphase (I, II) telophase (I, II) cytokinesis bivalent reductional division equational division G1 G2 S M G0 interphase n
c karyotype aneuploidy monsomic trisomic Down’s syndrome deletion duplication insertion inversion translocation non‐disjunction chromosome breakage polyploid x tetravalent endoreduplication endopolyploidy gene balance cellular network endosymbiont
organellarchromo
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STUDY QUESTIONS
2.1 Define chromatin. What is the difference between DNA, chromatin and chromosomes? 2.2 Species A has n=4 chromosomes and Species B has n=6 chromosomes. Can you tell from this information which species has more DNA? Can you tell which species has more genes? 2.3 The answer to question 2 implies that not all DNA within a chromosome encodes genes. Can you name any examples of chromosomal regions that contain relatively few genes?
2.4 a) How many centromeres does a typical chromosome have? b) What would happen if there was more than one centromere per chromosome? c) What if a chromosome had zero centromeres? 2.5 For a diploid with 2n=16 chromosomes, how many chromosomes and chromatids are per cell present in the gamete, and zygote and immediately following G1, S, G2, mitosis, and meiosis? 2.6 Bread wheat (Triticum aestivum) is a hexaploid. Using the nomenclature presented in class, an egg cell of wheat has
n=21 chromosomes. How many chromosomes in a zygote of bread wheat? 2.7 For a given gene: a) What is the maximum number of alleles that can exist in a 2n cell of a given diploid individual? b) What is the maximum number of alleles that can exist in a 1n cell of a tetraploid individual? c) What is the maximum number of alleles that can exist in a 2n cell of a tetraploid individual? d) What is the maximum number of alleles that can exist in a population? 2.8 a) Why is aneuploidy more often lethal than polyploidy? b) Which is more likely to disrupt gene balance: polyploidy or duplication?
2.9 For a diploid organism with 2n=4 chromosomes, draw a diagram of all of the possible configurations of chromosomes during normal anaphase I, with the maternally and paternally derived chromosomes labelled. 2.10 For a triploid organism with 2n=3x=6 chromosomes, draw a diagram of all of the possible configurations of chromosomes at anaphase I (it is not necessary label maternal and paternal chromosomes). 2.11 For a tetraploid organism with 2n=4x=8 chromosomes, draw all of the possible configurations of chromosomes during a normal metaphase.
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2%3*)!1($*3#(2#!/!01(<*#!,&/1,+!,-#4!	/1(#)!1()#:#()#(,!/()!2%3*)!.#!
1(-#&1,#)! 0#:/&/,#*46! ! I-10! 10! ,-#! ./010! %$!%#&'#"($! )*+$,! -!.+! /*0%!
2/**#)! /0#! -!.! 12! 345!"! 6#7+#7!,*1&+! 5-12-! 0,/,#0M! )3&1(<! </9#,#!
$%&9/,1%(+!,-#!,5%!9#9.#&0!%$!/!<#(#!:/1&!0#<&#</,#!$&%9!#/2-!%,-#&>!
N-/:,#&!O PABAIGNF!Q7!QBA!DQNRF
Figure 3.1 Pea plants
were used in the
discovery of some
fundamental laws of
genetics.
Q ( # ! D % 2 3 0 !S!8"9!!
!
#/2-! </9#,#! -/0! /(! #T3/*! :&%./.1*1,4! %$! 2%(,/1(1(<! #1,-#&!9#9.#&! %$!
,-#!<#(#!:/1&6!
!
!
!
'#()#*E0! 71&0,! D/5! 10! #0:#21/**4! 	/&;/.*#! .#2/30#! -#! 9/)#! -10!
%.0#&L/,1%(0!51,-%3,!;(%51(<!/.%3,!,-#!&#*/,1%(0-1:0!.#,5##(!<#(#0+!
2-&%9%0%9#0+!/()!CB@6!!J#!(%5!;(%5!,-/,!,-#!&#/0%(!,-/,!9%&#!,-/(!
%(#! /**#*#! %$! /! <#(#! 2/(! .#! :�#(,! 1(! /(! 1()1L1)3/*! 10! ,-/,! 9%0,!
#3;/&4%,12! %&</(1090! -/L#! /,! *#/0,! ,5%! 0#,0! %$! -%9%*%<%30!
2-&%9%0%9#06! !7%&!%&</(1090!,-/,!/&#!:&#)%91(/(,*4!)1:*%1)+!032-!/0!
-39/(0! %&! '#()#*E0! :#/0+! 2-&%9%0%9#0! #810,! 1(! :/1&0+! 51,-! %(#!
-%9%*%<!1(-#&1,#)!$&%9!#/2-!:/&#(,6! ! !C1:*%1)!2#**0!2/(!,-#&#$%&#!-%*)!
3:!,%!,5%!)1$$#&#(,!/**#*#0!%$!#/2-!<#(#+!51,-!%(#!/**#*#!%(!#/2-!9#9.#&!
%$! /! :/1&! %$! -%9%*%<%30! 2-&%9%0%9#06! ! G$! .%,-! /**#*#0! %$! /! :/&,123*/&!
<#(#! /&#! 1)#(,12/*+! ,-#! 1()1L1)3/*! 10! 0/1)! ,%! .#! 01:1;<715$! $%&! ,-/,!
<#(#6!!Q(!,-#!%,-#&!-/()+!1$!,-#!/**#*#0!/&#!)1$$#&#(,!$&%9!#/2-!%,-#&+!,-#!
<#(%,4:#! 10! 0#,#+1;<715$6! ! G(! 0%9#! 2/0#0+! 032-! /0! 5-#(! )#02&1.1(<!
<#(#0! %(! ,-#! 0#8! 2-&%9%0%9#0! %$! /! -39/(! 9/*#+! 5#! 30#! ,-#! ,#&9!
0#:*;<715$6! ! I-10! 10! .#2/30#! ,-#! 2#**! 2%(,/1(0! %(*4! %(#! U! /()! %(#! V!
2-&%9%0%9#+! 0%! ,-#&#! 10! %(*4! %(#! :%001.*#! /**#*#! $%&! #/2-! <#(#6!!
@*,-%3<-! /! ,4:12/*! )1:*%1)! 1()1L1)3/*! 2/(! -/L#! /,! 9%0,! ,5%! )1$$#&#(,!
/**#*#0! %$! /! :/&,123*/&! <#(#+! 9%&#! ,-/(! ,5%! /**#*#0! 2/(! #810,! 1(! /!
:%:3*/,1%(! %$! 1()1L1)3/*06! ! I-#! 9%0,! 2%99%(! /**#*#! 1(! /! (/,3&/*!
:%:3*/,1%(!10!2/**#)!,-#!.*"'",<=#!/**#*#6!!!!
HAD@IGQBFWGXF!"AIJAAB!@DDADAF!@BC!XWABQIVXAF!
@!0:#21$12!:%01,1%(!%(!/!2-&%9%0%9#!10!2/**#)!/!"1>5$6!!'%0,!%$!,-#!*%21!
<#(#,1210,0!)102300!/&#!%223:1#)!.4!<#(#0+!/()!,-#!,#&90!*%230!/()!<#(#!
/&#!%$,#(!30#)!1(,#&2-/(<#/.*46! !I-#!2%9:*#,#!0#,!%$!/**#*#0!/,!/(4!*%21!
%$!1(,#�,!1(!/(!1()1L1)3/*!)#$1(#!1,0!7#&1,<=#6!!I-#!)#,#2,/.*#!#$$#2,!%$!
,-#0#!/**#*#0!%(!,-#!0,&32,3&#!%&!$3(2,1%(!%$!,-/,!1()1L1)3/*!10!2/**#)!1,0!
Figure 3.2 Seven
traits Mendel studied
in peas.
Q ( # ! D % 2 3 0 !S!8"8!!
!
=0#&1,<=#6! ! ! I-#! :-#(%,4:#! 0,3)1#)! 1(! /(4! :/&,123*/&! <#(#,12!
#8:#&19#(,!9/4!&/(<#!$&%9!019:*#+!L101.*#!,&/1,0!032-!/0!-/1&!2%*%&+!,%!
9%&#!2%9:*#8!:-#(%,4:#0!1(2*3)1(<!)10#/0#!0302#:,1.1*1,4!%&!.#-/L1%&6!!!
!
D#,! 30! &#,3&(! ,%! /(! #8/9:*#! %$! /! 019:*#! :-#(%,4:#M! $*%5#&! 2%*%&! 1(!
'#()#*E0! :#/06!J#! -/L#! /*&#/)4! 0/1)! ,-/,! %(#! /**#*#! :&%)32#0! :3&:*#!
$*%5#&0+!5-1*#!,-#!%,-#&!/**#*#!:&%)32#0!5-1,#!$*%5#&0!Y71<3&#!O6OZ6!!"3,!
5-/,!)%#0! ,-10! ,#**!30!/.%3,! ,-#! $*%5#&!2%*%&!%$!/(! 1()1L1)3/*! ,-/,!-/0!
%(#! :3&:*#! /**#*#! /()! %(#! 5-1,#! /**#*#+! 1(! %,-#&! 5%&)0+! 5-/,! 10! ,-#!
=0#&1,<=#!%$!/(!1()1L1)3/*!5-%0#!<#(%,4:#!10!-#,#&%K4<%30?!J#!;(%5!
$&%9! #8:#&19#(,/*! )/,/! ,-/,! 1()1L1)3/*0! -#,#&%K4<%30! $%&! ,-#! :3&:*#!
/()!5-1,#! /**#*#0!%$! ,-#! $*%5#&!2%*%&! <#(#!-/L#!:3&:*#! $*%5#&06! ! ! ! I-#!
/**#*#!/00%21/,#)!51,-!:3&:*#!2%*%&!10!,-#&#$%&#!0/1)!,%!.#!'1:*&!&,!,%!
,-#! /**#*#! ,-/,! :&%)32#0! ,-#! 5-1,#! 2%*%&6! ! I-#! %,-#&! /**#*#+! 5-%0#!
:-#(%,4:#! 10!9/0;#)!.4!/!)%91(/(,!/**#*#! 1(!/!-#,#&%K4<%,#+! 10!2/**#)!
+#>#$$*?#6! ! ! Q$,#(+! /! )%91(/(,! /**#*#!51**! .#! &#:�#(,#)! .4! /! 2/:1,/*!
*#,,#&!Y#6<6!!Z!5-1*#!/!#001L#!/**#*#!51**!.#!&#:�#(,#)!1(!*%5#&!2/0#!
Y#6<6!"Z6!!W%5#L#&+!9/(4!)1$$#&#(,!040,#90!%$!<#(#,12!049.%*0!/&#!1(!30#!
YI/.*#! O6[Z+! 0%! 1,! 10! 19:%&,/(,! ,%! 3()#&0,/()! ,-#! )1$$#&#(,! ,4:#0! %$!
(%,/,1%(!/()!,%!30#!,-#9!2%(010,#(,*46!!!
#8/9:*#0!%$!
<#(%,4:#0!%$!
-#,#&%K4<%,#0!
,4:12/*!1(,#&:&#,/,1%(
!"! R::#&2/0#!*#,,#&Y0Z 9%0,!%$,#(!&#:�#(,0!)%91(/(,!/**#*#+!/()!
*%5#&2/0#!*#,,#&Y0Z!1()12/,#0!#001L#!/**#*#6!!!
"#"$%&!
!
D%5#&2/0#!*#,,#&0!51,-!\!03:#&02&1:,!&#:�#(,!51*)=,4:#!/**#*#+!
5-12-!10!%$,#(!)%91(/(,6!!@!*%5#&!2/0#!*#,,#&!51,-!%,-#&!
03:#&02&1:,0!Y%&!51,-!(%!03:#&02&1:,0Z!&#:�#(,0!%,-#&!/**#*#0+!
5-12-!/&#!%$,#(!#001L#6!
!'!&! I-#!(/9#!%$!,-#!*%230!10!1(!3::#&2/0#!%&!*%5#&!2/0#!*#,,#&0+!51,-!
03.02&1:,0!%&!03:#&02&1:,0!30#)!,%!1()12/,#!)1$$#&#(,!/**#*#06!!
P#(#&/**4+!(#1,-#&!/**#*#!10!2%9:*#,#*4!)%91(/(,+!%&!#*0#!,-#!
)%91(/(2#!&#*/,1%(0-1:0!/&#!3(;(%5(6!!
!
Figure 3.3
Relationship between
genotype and
phenotype for a
dominant allele.
Table 3.1 Examples of symbols used to represent alleles, and their dominance relationships
Q ( # ! D % 2 3 0 !S!8"@!!
!
"#01)#0! )%91(/(2#! /()! #001L1,4+! %,-#&! &#*/,1%(0-1:0! 2/(! #810,!
.#,5##(! /**#*#06! ! G(! $#:*"'1:*&!&>#! Y/*0%! 2/**#)! *&>1:="#,#!
'1:*&!&>#A!71<3&#!O6]Z+!.%,-!/**#*#0!/$$#2,!,-#!,&/1,!/))1,1L#*4+!/()!,-#!
:-#(%,4:#! %$! ,-#! -#,#&%K4<%,#! 10! 1(,#&9#)1/,#! .#,5##(! #1,-#&! %$! ,-#!
-%9%K4<%,#06! ! 7%&! #8/9:*#+! /**#*#0! %$! $%&! &#)! 2%*%&! 1(! 2/&(/,1%(0! /()!
0%9#!%,-#&!0:#21#0!#8-1.1,!0#91=)%91(/(2#+!0%!,-/,!/!-#,#&%K4<%,#!-/0!
:1(;!:#,/*0+!5-1*#! /! -%9%K4<%,#! $%&! %(#! /**#*#! -/0! &#)!:#,/*0! /()! ,-#!
%,-#&! -/0! 5-1,#! :#,/*06! ! ! I-30! 1(! 0#91=)%91(/(2#+! ,-#! )%0/<#! %$! ,-#!
/**#*#0!/$$#2,0!,-#!:-#(%,4:#+!#6<6! ,-#!:-#(%,4:#!%$!,-#!-#,#&%K4<%,#!10!
/::&%819/,#*4!-/*$!/0!0,&%(<!/0!,-#!:-#(%,4:#!%$!/!-%9%K4<%,#6!!!!!!
!
!
B1"'1:*&!&>#! 10! /(%,-#&! ,4:#! %$! /**#*12! &#*/,1%(0-1:+! 1(! 5-12-! /!
-#,#&%K4<%30! 1()1L1)3/*! #8:�#0! ,-#! :-#(%,4:#! %$! .%,-! /**#*#0!
0193*,/(#%30*46! !@(!#8/9:*#!%$!2%=)%91(/(2#!10!$%3()!51,-1(!,-#!@"Q!
.*%%)!<&%3:!%$!-39/(0+!5-12-! 10!2%(,&%**#)!.4!/!<#(#!2/**#)!(!Y71<3&#!
O6^Z6! ! I-#&#! /&#! ,-&##! :%001.*#! /**#*#0! /,! ,-10! *%230M! (!+! ()+! /()! *6!!!
W%9%K4<%30!1()1L1)3/*0!$%&!(!!%&!()!:&%)32#!%(*4!@!%&!"!,4:#!/(,1<#(0+!
�:#2,1L#*4+! %(! ,-#! 03&$/2#! %$! ,-#1&! .*%%)! 2#**0+! /()! ,-#&#$%&#! -/L#!
#1,-#&!,4:#!@!%&!,4:#!"!.*%%)6!!W#,#&%K4<%30!(!()!1()1L1)3/*0!-/L#!.%,-!
@! /()! "! /(,1<#(0! %(! ,-#1&! 2#**! 03&$/2#+! /()! 0%! -/L#! ,4:#! @"! .*%%)6!!
B%,12#!,-/,!,-#!-#,#&%K4<%,#!#8:�#0!.%,-!/**#*#0!0193*,/(#%30*4+!/()!
10!(%,!0%9#!;1()!%$!(%L#*!1(,#&9#)1/,#!.#,5##(!@!/()!"6!!N%=)%91(/(2#!
10! ,-#&#$%&#! )10,1(2,! $&%9! 0#91=)%91(/(2#+! /*,-%3<-! ,-#4! /&#!
0%9#,19#0!2%($30#)6!!!'/(4!,4:#0!%$!9%*#23*/&!9/&;#&0+!5-12-!5#!51**!
)102300! 1(! /! */,#&! 2-/:,#&+! )10:*/4! /! 2%=)%91(/(,! &#*/,1%(0-1:! /9%(<!
/**#*#06!!G,!10!/*0%!19:%&,/(,!,%!(%,#!,-/,!,-#!,-1&)!/**#*#+!*+!)%#0!(%,!9/;#!
/(4! /(,1<#(0! /()! 10! #001L#! ,%! /**! %,-#&! /**#*#06! ! ! W%9%K4<%30!
#001L#! 1()1L1)3/*0! Y**Z!-/L#! ,4:#!Q!.*%%)6!I-10! 10! /!30#$3*! [()#&!
,-/,! )1$$#&#(,! ,4:#0! %$! )%91(/(2#! &#*/,1%(0-1:0! 2/(! #810,+! #L#(! $%&!
/**#*#0!%$!,-#!0/9#!<#(#6!
Figure 3.4
Relationship between
genotype and
phenotype for semi-
dominant alleles.
Q ( # ! D % 2 3 0 !S!8"C!!
!
!
"GQNWA'GN@D!"@FGF!Q7!CQ'GB@BNA!
J#!2/((%,!:&#)12,!5-#,-#&!/(!/**#*#!51**!.#!)%91(/(,!./0#)!019:*4!%(!
,-#!:-#(%,4:#!%$!-%9%K4<%,#0>!5#!930,!/*0%!;(%5!,-#!:-#(%,4:#!%$!/!
-#,#&%K4<%,#6! ! J-/,+! ,-#(+! 2/30#0! )%91(/(2#?! ! I-#&#! /&#! 9/(4!
)1$$#&#(,!.1%2-#912/*!2%()1,1%(0!,-/,!9/4!9/;#!%(#!/**#*#!)%91(/(,!,%!
/(%,-#&+!.3,!%(#!%$!,-#!9%0,!2%99%(!10!0!="1$522*>*#&><6!!'%0,!<#(#0!
:&%)32#! 9%&#! ,-/(! #(%3<-! :&%,#1(! ,%! /22%9:*10-! ,-#1&! (%&9/*!
.1%*%<12/*!$3(2,1%(6!!G$!/(!/**#*#!10!-/:*%03$$121#(,+!,-#(!_30,!%(#!2%:4!%$!
,-/,! /**#*#! :&%)32#0! #(%3<-! :&%,#1(! ,%! -/L#! ,-#! 0/9#! #$$#2,! /0! ,-#!
/9%3(,! %$! :&%,#1(! :&%)32#)! .4! ,5%! 2%:1#0! %$! ,-/,! /**#*#6! ! 7%&! ,-#!
9/_%&1,4! %$! <#(#0! 0,3)1#)+! ,-#! (%&9/*! Y16#6! .*"'",<=#Z! /**#*#0! /&#!
-/:*%03$$121#(,+!0%!#L#(!1$!/!93,/,1%(!2/30#0!/!2%9:*#,#!*%00!%$!$3(2,1%(!
1(! %(#! /**#*#+! ,-#! 51*)=,4:#! /**#*#! 51**! .#! )%91(/(,! /()! ,-#! (%&9/*!
$3(2,1%(! %$! ,-#! .1%2-#912/*! :/,-5/4!51**! .#! &#,/1(#)6! ! ! Q(! ,-#! %,-#&!
-/()+! 1(! 0%9#!.1%2-#912/*!:/,-5/40+! /! 01(<*#!51*)=,4:#!/**#*#!9/4!.#!
0!="1*&$522*>*#&,+!16#6!1,!9/4!(%,!:&%)32#!#(%3<-!:&%,#1(!,%!*,!1(!/!
(%&9/*!:-#(%,4:#!1$!,-#!%,-#&!/**#*#!1(!/!-#,#&%K4<%,#!10!(%,!$3(2,1%(/*6!!
G(! ,-10! 2/0#+! /! (%(=$3(2,1%(/*! /**#*#! 51**! .#! )%91(/(,! ,%! /! 51*)=,4:#!
/**#*#6!
IWA!XRBBAII!F`R@HA!@BC!'QBQWV"HGC!NHQFFAF!
P#(#,1210,0+!1(2*3)1(<!'#()#*+!9/;#!30#!%$!,+5#!D+##'*&7!"*&#$!Y71<3&#!
O6a!/Z6!!I-#0#!/&#!:%:3*/,1%(0!%$!:*/(,0!%&!/(19/*0!1(!5-12-!/**!:/&#(,0!
/()! ,-#1&! %$$0:&1(<! -/L#! 1)#(,12/*! :-#(%,4:#0! %L#&!9/(4! <#(#&/,1%(0!
51,-! �:#2,! ,%! /! :/&,123*/&! ,&/1,6! I&3#! .&##)1(<! *1(#0! /&#! 30#$3*+!
.#2/30#! ,-#4! 2/(! .#! /0039#)! ,%! .#! -%9%K4<%30! $%&! ,-#! /**#*#0! ,-/,!
/$$#2,!/!,&/1,!%$!1(,#�,6!J-#(!,5%!1()1L1)3/*0!,-/,!/&#!-%9%K4<%30!$%&!
,-#! 0/9#! /**#*#0! /&#! 2&%00#)+! /**! %$! ,-#1&! %$$0:&1(<! 51**! /**! /*0%! .#!
-%9%K4<%306!
Figure 3.5
Relationship between
genotype and
phenotype for co-
dominant alleles (IA,
IB), and a recessive
allele.
Q ( # ! D % 2 3 0 !S!8"E!!
!
!
!
P1L#(! ,-#! <#(%,4:#0! %$! /(4! ,5%! :/&#(,0+! 5#! 2/(! :&#)12,! /**! %$! ,-#!
:%001.*#!<#(%,4:#0!%$!,-#!%$$0:&1(<6! !73&,-#&9%&#+! 1$!5#!/*0%!;(%5!,-#!
)%91(/(2#! &#*/,1%(0-1:0! $%&! /**! %$! ,-#! /**#*#0+! 5#! 2/(! :&#)12,! ,-#!
:-#(%,4:#0!%$! ,-#!%$$0:&1(<6! !@! 2%(L#(1#(,!9#,-%)! $%&! 2/*23*/,1(<! ,-#!
#8:#2,#)! <#(%,4:12! /()! :-#(%,4:12! &/,1%0! $&%9! /! 2&%00! 10! ,-#! 30#! %$!
X3((#,,!FT3/&#+!5-12-!10!/!9/,&18!1(!5-12-!/**!%$!,-#!:%001.*#!</9#,#0!
:&%)32#)!.4!%(#!:/&#(,!/&#!*10,#)!/*%(<!%(#!/810+!/()!,-#!</9#,#0!$&%9!
,-#! %,-#&! :/&#(,! /&#! *10,#)! /*%(<! ,-#! %,-#&! /8106! ! A/2-! :%001.*#!
2%9.1(/,1%(! %$! </9#,#0! 10! *10,#)! /,! ,-#! 1(,#&0#2,1%(! %$! #/2-! &%5! /()!
2%*39(6!!!
J-#(! ,5%! -#,#&%K4<%,#0! /&#! 2&%00#)+! 5-/,! <#(%,4:#0! /&#! :&%)32#)+!
/()!5-/,! 10! ,-#!#8:#2,#)! $&#T3#(24!%$! #/2-!<#(%,4:#?! ! ! G$!5#!30#! ,-#!
049.%*0!!!/()!"!,%!&#:�#(,!#/2-!,-#!,5%!/**#*#0!%$!,-#!-#,#&%K4<%,#+!
,-#(!$&%9!,-#!F5&&#,,!645!+#!Y71<3&#!O6bZ!5#!0##!,-/,!,-&##!<#(%,4:#0!
/&#! :&%)32#)! 1(! /! &/,1%! %$! [McM[6! ! G$! 5#! ;(%5! 0%9#,-1(<! /.%3,! ,-#!
:-#(%,4:#0!/()!)%91(/(2#!&#*/,1%(0-1:0!%$!,-#0#!/**#*#0!Y/0!19:*1#)!.4!
,-#!049.%*0!30#)Z+!5#!2/(!:&#)12,!,-#!#8:#2,#)!:-#(%,4:12!&/,1%!%$!,-#!
:&%<#(4! $&%9! ,-10! 2&%00! 10! OM[6! ! @! 2&%00! .#,5##(! ,5%! 1()1L1)3/*0! ,-/,!
/&#!.%,-!-#,#&%K4<%30!/,!/! 01(<*#! *%230! 10! 0%!2%99%(! 1(!<#(#,120! ,-/,!
,-10!2&%00!10!<1L#(!1,0!%5(!(/9#M!!,-#!:1&10<D+*'!>+1$$!Y71<3&#!O6a!.Z6!!!
! !! "
!! !! !"
"! !" ""
!
!
Figure 3.6 a) a true-
breeding line b) a
monohybrid cross
produced by mating
two pure-breeding
lines
Figure 3.7 A Punnett
Square showing a
monohybrid cross
Q ( # ! D % 2 3 0 !S!8"G!!
!
IAFI!NHQFFAF!N@B!"A!RFAC!IQ!CAIAH'GBA!PABQIVXAF!
d(%51(<!,-#!<#(%,4:#0!%$!/(!1()1L1)3/*!10!303/**4!/(!19:%&,/(,!:/&,!%$!
/!<#(#,12!#8:#&19#(,6!!W%5#L#&+!<#(%,4:#0!2/((%,!.#!%.0#&L#)!)1,*4>!
,-#4!930,!.#!1($#&&#)!./0#)!%(!:-#(%,4:#06!"#2/30#!%$!)%91(/(2#+!1,!10!
%$,#(! (%,! :%001.*#! ,%! )10,1(<310-! .#,5##(! /! -#,#&%K4<%,#! /()! /!
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Figure 3.7 Reciprocal
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sex-linked trait.
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Figure 3.9 Probability
distribution of the chi-
square statistic for five
different degrees of
freedom
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A1A1 A1A2 A2A2
1 all hairs black on the same
individual:
50% of hairs are all
black
50% of hairs are all
white
all hairs white
2 all hairs black all hairs are the same
shade of grey
all hairs white
3 all hairs black all hairs black 50% of individuals
have all white hairs
50% of individuals
have all black hairs
4 all hairs black all hairs black mice have no hair
5 all hairs black all hairs white all hairs white
6 all hairs black all hairs black all hairs white
7 all hairs black all hairs black hairs are a wide range
of shades of grey
!
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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.
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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.
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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 pregnant woman resulted in birth defects. Finally ethidium bromide, the dye that fluorescently stains DNA in laboratory assays, is 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
Figure 4.6 Alkylation of guanine (shown in red) allows G to bond with thymine rather than cytosine at replication.
Figure 4.7 Benzopyrene (circled in red) is an example of an intercalating agent
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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 lossoffunction 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 gainoffunction, producing either more of the active protein (hypermorph) or producing an active protein with a new function (neomorph).
Figure 4.8 Thymine dimers are formed when adjacent thymine bases on the same DNA strand become covalently linked (red bonds) follow exposure to mutagens such as UV light. The dimers distort base pairing and can interrupt processes such as replication.
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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.
Some phenotypes require individuals to reach a particular developmental stage before they can be scored. For example, flower color can only be scored in plants that are mature enough to make flowers, and eye color can only be scored in flies that have developed eyes. However, some alleles may not develop sufficiently to be included among the progeny that are scored according to their phenotype. Lethal alleles that arrest the development of an individual at an embryonic stage may therefore go unnoticed in a typical mutant screen. Furthermore, the progeny of a monohybrid cross involving an embryonic lethal recessive allele may therefore all be of a single
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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
Figure 4.9 In this simplified biochemical pathway, two enzymes encoded by two different genes modify chemical compounds in two sequential reactions to produce a purple pigment. Loss of either of the enzymes disrupts the pathway and no pigment is produced.
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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.
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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.
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.
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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 mutant polymorphism insertion deletion substitution mutagen biological mutagen chemical mutagen physical mutagen tautomer mispairing loop SSR insertional mutagen Class I, Class II
transposon retrotransposon reverse transcriptase transposase non‐autonomous autonomous SINE, LINE, Alu P‐element T‐DNA alkylation agent EMS intercalating agent benzopyrene ethidium bromide thymine dimer mutant screen
loss‐of‐function gain‐of‐function amorph null hypomorph hypermorph neopmorph somatic germline M0, M1, M2 silent mutation redundancy lethality complementation group
_______________________________________________________________________________________________________________
STUDY QUESTIONS
4.1 How are polymorphisms and mutations alike? How are they different? 4.2 What are all of the ways a substitution can occur in a DNA sequence? 4.3 What are all of the ways a deletion can occur in a DNA sequence? 4.4 What are all of the ways an insertion can occur in a DNA sequence? 4.5 In the context of this chapter, explain the health hazards of smoking tobacco. 4.6 You have exposed a female fruit fly to a mutagen. Mating this fly with a non‐mutagenized male produces offspring that appear to be completely normal.
However there are twice as many females as males in the F1 progeny of this cross. a) Propose a hypothesis to explain these observations. b) How could you test your hypothesis? 4.7 You decide to use genetics to investigate how your favourite plant makes its flowers smell good. a) What steps will you take to identify some genes that are required for production of the sweet floral scent? Assume that this plant is a self‐pollinating diploid. b) One of the recessive mutants you identified has fishy‐smelling flowers, so you name the mutant (and the mutated gene) fishy. What do you hypothesize about the normal function of the wild‐type fishy gene?
c) Another recessive mutant lacks floral scent altogether, so you call it nosmell. What could you hypothesize about the normal function of this gene? 4.8 Suppose you are only interested in finding dominant mutations that affect floral scent. a) What do you expect to be the relative frequency of dominant mutations, as compared to recessive mutations, and why? b) How will you design your screen differently than in the previous question, in order to detect dominant mutations specifically? c) Which kind of mutagen is most likely to produce dominant mutations, a mutagen that produces point mutations, or a mutagen that produces large deletions? 4.9 Which types of transposable elements are transcribed?
4.10 You are interested in finding genes involved in synthesis of proline (Pro), an amino acid that is normally synthesizes by a particular model organism. a) How would you design a mutant screen to identify genes required for Pro synthesis? b) Imagine that your screen identified ten mutants (#1 through #4) that grew poorly unless supplemented with Pro. How could you determine the number of different genes represented by these mutants? c) If each of the four mutants represents a different gene, what will be the phenotype of the F1 progeny if any pair of the four mutants are crossed? d) If each of the four mutants represents the same gene, what will be the phenotype of the F1 progeny if any pair of the four mutants are crossed?
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Figure 5.1 Polydactyly
is an example of a
human trait that can be
studied by pedigree
analysis
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Figure 5.2 Symbols
used in drawing a
pedigree.
Figure 5.3 A pedigree
consistent with AD
inheritance.
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Figure 5.4 Two
pedigrees consistent with
XD inheritance.
Figure 5.5 Some types
of rickets may follow an
XD mode of inheritance.
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Figure 5.8 A pedigree consistent with XR inheritance. Figure 5.9 Some forms of colour
blindness are inherited as XR-traits.
Colour blindness is diagnosed using tests
such as this Ishihara Test.
Figure 5.10
Stephen Hawking
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Figure 5.11
Individuals in this
pedigree are labeled
with numbers to make
discussion easier
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Figure 5.12
Allele frequencies may
also be studied on the
population level.
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Photo and Illustration Credits
Ch Fig source author license1 1 0 flickr eclectic echoes CC: AND
1 1 original unknown PD
1 2 original Deyholos CC: AN
1 3
J. Exp. Med. 98:21, 1953 Austrian, Robert pending
1 4 original Deyholos CC: AN
1 5 Wikipedia Colm, Graham PD
1 6 Wikipedia (modified) Adenosine, Deyholos CC: AS
1 7 Wikipedia Ströck, Michael GFDL
1 8 original Deyholos CC: AN
1 9 original Deyholos CC: AN
1 10 flickr Westby, Max CC: ANS
1 10 flickr Joly, David CC: ANS
1 10 Wikipedia Altun, Zeynep F. CC: AS
1 10 Wikipedia Masur GFDL
1 10 Wikipedia Azul GFDL
1 10 Wikipedia unknown GFDL
2 1 flickr: TheJCB
Zhang et al. (2007) J. Cell Biol. 177:231-242. CC: ANS
2 2 Wikipedia Wheeler, Richard GFDL
2 3 unknown unknown pending
2 4 original Deyholos CC: AN
2 5 BMC unknown pending
2 6 PLoS Genetics
Somma MP et al. (2008) PLoS Genets 4(7): e1000126 PD
2 7 original Deyholos CC: AN
2 8 PLoS Genetics
Chelysheva, L. et al (2008) PLoS Genetics PD
2 9 original Deyholos CC: AN
2 10 original Deyholos CC: AN
2 11 Wikipedia NHGRI PD
2 12 Wikipedia Zephyris GFDL
2 13 flickr Bell, Darwin CC: AN
2 14 flickr Elissa Lei, Ph.D. @ NIH CC: A
3 1 flickr Guthier, Christian CC: A
3 2 Wikipedia Ruiz, Mariana PD
3 3 original Deyholos (Fireworks) CC: AN
3 4 original Deyholos CC: AN
3 5 original Deyholos CC: AN
3 6 original Deyholos CC: AN
3 7 original Deyholos CC: AN
3 8 original Deyholos CC: AN
3 9 Wikipedia PAR PD
4 1 flickr ecstaticist CC: ANS
4 2 PLoS Biology
Zarbalis, K. et al (2004) PLoS Biology PD
4 3 original Deyholos CC: AN
4 4 original Deyholos CC: AN
4 5 original Deyholos CC: AN
4 6 original Deyholos CC: AN
4 7 Wikipedia Zephyris GFDL
4 8 original Deyholos CC: AN
4 9 original Deyholos CC: AN
4 10 original Deyholos CC: AN
4 11 original Deyholos CC: AN
4 B1 original Deyholos CC: AN
4 B2 Wikipedia unknown PD
4 B2 flickr windy234 CC: AN
5 1 Wikipedia en:User:Drgnu23 GFDL
5 2 original Deyholos CC: AN
5 3 original Deyholos CC: AN
5 4 original Deyholos CC: AN
5 5 Wikipedia Mrich CC: AS
5 6 original Deyholos CC: AN
5 7 Wikipedia
U.S. Air Force photo/Staff Sgt Eric T. Sheler PD
5 8 original Deyholos CC: AN
5 9 Wikipedia unknown PD
5 10 Wikipedia NASA PD
5 11 original Deyholos (Fireworks) CC: AN
5 12 flickr Stern, Zach CC: AND
6 1 flickr Gossamer1013 CC: AND
6 2 original Deyholos CC: AN
6 3 original Deyholos CC: AN
6 4 original Deyholos CC: AN
6 5 flickr Curley, John CC: AN
6 5 flickr Romans, Phil CC: AND
6 5 flickr Miss Chien CC: AND
6 6 original Deyholos CC: AN
6 7 flickr unknown CC: AD
6 8 original Deyholos CC: AN
6 9 original Deyholos CC: AN
6 10 original Deyholos CC: AN
6 11 other UN Lincoln pending
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6 13 original Deyholos CC: AN
7 1 Wikipedia Abiyoyo CC: AS
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7 3 original Deyholos CC: AN
7 4 Wikipedia Morgan PD
7 5 original Deyholos CC: AN
7 6 original Deyholos CC: AN
7 7 original Deyholos CC: AN
7 8 original Deyholos CC: AN
7 9 original Deyholos CC: AN
7 10 NCBI NIH PD
7 11 original Deyholos CC: AN
7 12 original Deyholos CC: AN
7 13 original Deyholos CC: AN
8 1 flickr estherase CC: ANS
8 2 original Deyholos CC: AN
8 3 original Deyholos CC: AN
8 4 Wikipedia madprime GFDL
8 5 original Deyholos CC: AN
8 6 Wikipedia madprime GFDL
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8 8 original Deyholos CC: AN
8 9 original Deyholos CC: AN
8 10 flickr DeathByBokeh CC: AN
8 11 flickr 457088634_585df11af5_o pending
8 12 Wikipedia Magnus Manske PD
8 13 Wikipedia Transcontrol GFDL
8 14 original Deyholos CC: AN
9 1 flickr Jaime Golombek CC: AND
9 2 original Deyholos CC: AN
9 3 original Deyholos CC: AN
9 4 original Deyholos CC: AN
9 5 original Deyholos CC: AN
9 6 original Deyholos CC: AN
9 7 original Deyholos CC: AN
9 9 original Deyholos CC: AN
9 10 original Deyholos CC: AN
10 1 Goodsell, Scripps Goodsell, Scripps EDU
10 2 original Deyholos CC: AN
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10 6 Wikipedia Abizar Lakdawalla PD
10 7 NCBI unknown PD
10 8 original Deyholos CC: AN
11 1 flickr frenquency CC: AND
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11 5 original Deyholos CC: AN
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11 7 original Deyholos CC: AN
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11 9 original Deyholos CC: AN
11 10 original Deyholos CC: AN
11 11 flickr Beardy Git CC: AND
11 12 original Deyholos CC: AN
12 1 flickr Uthman, Ed CC: AS
12 2 Wikipedia NIH PD
12 3 Wikipedia Hayman, J PD
12 4 Wikipedia unknown PD
12 5 flickr Uthman, Ed CC: AS
12 6 Wikipedia Mark 'AbsturZ' PD
12 7 Wikipedia
Splettstoesser, Thomas based on Cho et al. Science 265 pp. 346, 1994 CC: AS
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