Objectives Unit Twelve:  Mutation and Mutagenesis

1.  Define the following terms and utilize these terms in an appropriate context:
        acridine dye          alkylating agent         base analogue             DNA ligase
        DNA polymerase excision repair           frameshift mutation      induced mutation
        lethal mutation      mismatch repair        nutritional mutation       photoreactivation enzyme
        point mutation       regulatory mutation   tautomeric shift             temp-sensitive mutation
        transposon

2.  Distinguish mutations according to tissues being affected, somatic mutations versus
    gametic mutations.

3.  Relate the type of mutation to the effects on the individual as well as those effecting gene
    frequencies and the gene pool.

4. Classify mutations according to effects on the person receiving such genes as
    morphological, biochemical, lethal, conditional and regulatory.

5.  Differentiate between a chromosomal mutation and a gene mutation.

6.  Explain why a random mutation is more likely to be deleterious than beneficial?

7.  Explain why mutations in a diploid organism are usually recessive.

8.  Describe the occurance of spontaneous mutations that arise naturally as a result of chemical
     rearrangements, tautomeric shifts or as the result of errors occurring during DNA replication.

9.  Describe tautomerism and the way in which this chemical event may lead to mutation.

10.  Describe and illustrate a point mutation and its effects on a gene sequence.

11.  Describe how base analogues and alkylating agents and tautomeric shifts cause chemical
    changes in nucleotides leading to base-pairing problems resulting in base substitution
    mutations (point mutations) arising following DNA replication.

12.  Describe and illustrate a frame shift mutation and its effects on a gene sequence.

13.  Catagorize missense and nonsense modifications of a gene sequence.

14.  Describe the effect of agents ,such as radiation and acridine dyes, on a gene sequence
    with relation to their ability to cause frame shift mutations.

15.  Explain why frameshift mutations are likely to be more detrimental than a point mutation.

16.  Contrast the various types of DNA repair mechanisms known to counteract the effects of
        UV light and other DNA damage.

17.  Differentiate between excision repair and photoreactivation mechanisms of repair.

18.  If given a short wild-type protein sequence and several mutated versions of this sequence:
    -be able to identify likely mRNA codons relating to the given protein
    -be able to identify the specific and most likely mutational event that created the new mutated
    sequence.

Resources:  Text Chapter 14,  Cartoon Guide pgs. 79-83; 158-163; 179-184
 

Mutation and Mutagenesis

Mutation- failure to store genetic info faithfully; change in stored info
   reflected in expression

    Mutation includes chromosomal changes and changes within single genes

    *Gene mutation- Hugo DeVries 1901     Attention to: classification, origin, induction

    Genetics would not be if it weren't for mutation

    Classification of Mutations
        Various schemes, not mutually exclusive
        -depend on which aspect of mutation being investigated

            1. Spontaneous vs. Induced Mutation
                All mutations either:
                    A. spontaneous- no specific agent other than natural forces

                        -arise randomly, normal chemical processes, background radiation, alterations
                            in chemical structure

                    B. Induced- result of influence of an artificial factor
                        -X-rays, wide range of chemical mutagens

            2. Gametic vs. Somatic mutations
                    A. Gametic- mutation in sex cell (germ-line, germinal mutations)
                        - hemophilia perhaps best known (sex-linked recessive)

                    B.Somatic- non sex cells

                        -changes not passed on to future generations
                        -mutation in single cell may not impare the organism even if mutation is
                            detrimental

                        a. mutation may occur in cell not active or essential
                        b. even if critical gene affected, thousands of others remain

                        -greatest impact; if dominant and occur early in devel.
                        -autosomal recessives require 2 genes for expression

            3. Classification on basis of their effect on organism

                    A. morphological mutation (visible)
                        -variations deviate from normal or wild-type phenotype

                    B. Nutritional or biochemical variations
                        -bacteria, fungi (AMP+), synthesis of amino acids or vitamins

                        -Hemophilia, sickle cell anemia
                        *not visible and do not usually affect specific morphological characters;
                            more general effect on well being

                    C. Behavioral mutations
                        -affect behavior patterns of an organism (mating behavior, circadian rhythms)

                        -often difficult to discern
                        Ex. mating behavior of fruit fly impared because it can't beat its wings

                    ??flight muscle, nerve ending, brain???

                    D. Regulatory mutations
                        -may disrupt transcription, permanently activate or inactivate a gene

                    E. Lethal mutations
                        -Huntingtons disease, Tay Sachs
                        -severity does not allow individual to survive

                        -inactivating gene for RNA polymerase
                      Haploids die immediately- no wild type gene to mask

                      Diploids can carry lethal if masked by dominant  (most are recessive, however
                        Huntingtons dom.)

                    F. Conditional mutations
                        -affect mutant under certain conditions

                    Ex. Temperature sensitive mutation
                        -Permissive temps: mutant gene functions normally
                        -Restrictive temps: lose functional ability

Effects of Mutation on DNA

1. Point mutation
        A. missense- alters base creating a change of an amino acid (missense because not
            the same original message)

        B. nonsense- original codon converted to termination codon

            -simplest change is single base (nucleotide)
                Transition- purine replaces purine or pyrimidine replaces pyrimidine

                Transversion- purine replaces pyrimidine or visa versa (*more damaging)

                    Changes in third pair of codon:
                        if pur to pur or pyrm to pyrm- likely to code for same amino acid

                        if purine to pyrimidine- likely to be different AA

2. Frame shift Mutation

    -insertion or deletion of single nucleotide at any point in gene

    -severe, cause change in reading frame after mutation
 

                                                    THE CAT SAW THE DOG

                           POINT MUTATION                                 FRAME SHIFT MUTATION
                            THE BAT SAW THE DOG                       Gain: THE CAT SAW THE ZDO G
                            THE CAT SAW THE HOG                            (addition of Z)
                            THE CAT SAT THE DOG                                   THE CMA TSA WTH EDO G
                                                                                                      (addition of M)
                            *missense
                                                                                                 Loss: THE CAT SAW TED OD
                                                                                                      (loss of original H)
                                                                                                            THE ATS AWT HED OG
                                                                                                      (loss of original C)

                                                                                                    *Nonsense

Molecular basis of Mutation (Chemical)
    1. Replication machinery
            Mutators- proofreading function lost (alteration of DNA poly III)

    2. Tautomeric Shift (Tautomerization)
        -bases of DNA occur in 2 or more forms, differ in proton shift
        -Result in base pair changes (Point mutation)

        -Biologically important tautomers involve:
                Keto-enol pairs for thymine and guanine

                Amino-imino pairs for adenine and cytosine

         *infrequent tautomer capable of bonding with normally  noncomplementary base.
            However, pairing still purine/pyrimidine

                End result A=T becomes G=C, of course next round of replication incorporates
                the mutation

                *A ordinarily pairs w/ T, in its rare tautomeric form  A pairs w/ C

    3. Deamination- loss of amino group

        Cytosine --> deaminated --> carbonyl replaces amino and converts cytosine to uracil,
            therefore, pairs with A not G

        *usually corrected by polymerase because U not usually component of DNA

            5-methylcytosine - common modified base, pairs w/ G deaminated form pairs w/ T;
                more serious because T is normal component of DNA, therefore, not corrected
                "Hot spots"

    4. Base analogues
        -mutagenic molecules that can substitute for purines or  pyrimidines during nucleotide
            and DNA synthesis

        -resembles Thymine

        -causes A=T to G=C transition

    5. Nitrous Acid, Hydroxylamine

    6. Alkylating Agents
        electrophilic- attracted to negative charges

        DNA- each nucleotide contains full and partial negative charges

        Electrophiles attack and add alkyl groups (CH3 or CH2-CH3)

        Mustard Gases, most environmental carcinogens (breaks in nucleotides, mispairing)

    7. Acridine Dyes ---> frame shifts
        -about same dimension as nitrogenous bases

        -intercalate, wedge between purines and pyrimidines of intact DNA. Cause additions
            and deletions

        Proflavin, acridine orange
        -mispairing- looping

Radiation induced mutation

    High energy- Gamma radiation, X-ray (ionizing radiation)

    U.V.- shorter wavelength than visible light

    Ultraviolet light
        -purines and pyrimidines absorb UV most intensly at 260nm
        -major effect on pyrimidines; addition of H2O to ring

            1. pyrimidine dimers formed, usually between 2 thymines
            2. dimers distort DNA conformation and inhibit normal  replication

                -UV ordinarily not strong enough to penetrate layers of skin

                -Xeroderma pigmentosum- autosomal recessive in humans, predisposes
                    individuals to epidermal pigment abnormalities

                    1. exposure to sunlight results in malignant growths on skin
 

High Energy- Gamma, X-ray, cosmic rays (very short wavelength)
    -strong enough to penetrate deeply into tissues causing  ionization of molecules
        encountered along the way.

    -as X-ray penetrates, stable molecules become free radicals  trail of ions can initiate
        chemical reactions that can directly or indirectly effect genetic material

    -frequently causes single or double strand breaks

    -linear relationship between radiation dose and induction of mutation
        *each doubling of dose produces doubling of mutations

           General observations

            -intensity of the dose administered seems to make little difference in mutagenic rate.

             Ex. 2000-roentgen exposure, either in a single dose or cumulative w/many smaller
                doses produces same effect

            (1 roentgen- 2 billion ionizations/cm3)

            -certain portions of cell cycle more susceptible
                X-rays---> breaks, deletions, translocations, chromosome fragmentation damage
                    occurs most readily  when chromosomes are greatly condensed:  Mitosis

            *also reasoning behind its use as a cancer treatment

DNA Repair

    1. Reversion- repair through mutation pg 286-287
    2. Photoreactivation-
        -UV induced damage could be partially reversed if cells were briefly exposed to light in
            blue range.

        -temp dependent, involves enzymatically controlled mechanism

        Photoreactivation enzyme (PRE)- cleaves bonds between thymine dimers.  Enzyme
            must absorb a photon of light to cleave dimer.

    3. Excision repair- activated in response to any DNA damage thatdistorts helix
        -apurinic or apyrimidinic site where base removed from sugar
        -endonuclease recognizes damage and clips out DNA

        -DNA polymerase fills gap; DNA ligase seals final nick

    4. Proofreading and mismatch repair
        -spontaneous errors occur ~ 1/100,000 ; immediately subject to proofreading.
            Thought to increase fidelity to 1/1,000,000

            Mismatch repair
                1. mismatch must be detected
                2. incorrect nucleotide removed
                3. replacement w/correct base

                *special problem:  no distortion of the helix to signal that problem exists like UV

                *if detected how is the mismatched base discriminated from the "correct"

            DNA Methylation
                -methyl group (CH3) is added to adenine on parent DNA
                -newly synthesized DNA remains temporarily unmethylated, therefore, new can be
                    discerned from old

                -methyl added to adenines in sequence 5'GATC 3'
                                                                                     3'CTAG 5'
                     these occur every 250 bases, usually not far away

    5. Recombinational Repair- responds when damaged DNA has escaped repair
        -As DNA replicated polymerase stalls and then skips damage creating gap
        -protein directs recombination; segment homologous to damaged segment inserted into
            gap
        -resulting gap on donor strand repaired by polymerase and ligase action
 

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