The Molecular Basis of Inheritance

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1 hapter 6 he Molecular Basis of Inheritance Dr. Wendy Sera Key oncepts in hapter 6. DN is the genetic material. Many proteins work together in DN replication and repair. 3. chromosome consists of a DN molecule packed together with proteins. Houston ommunity ollege Biology 406 Life s Operating Instructions In 953, James Watson and Francis rick introduced an elegant double-helical model for the structure of deoxyribonucleic acid, or DN Life s Operating Instructions, cont. Hereditary information is encoded in DN and reproduced in all cells of the body his DN program directs the development of biochemical, anatomical, physiological, and (to some extent) behavioral traits DN is copied during DN replication, and cells can also repair their DN Figure 6. What is the structure of DN? oncept 6.: DN is the genetic material Early in the 0th century, the identification of the molecules of inheritance loomed as a major challenge to biologists he Search for the enetic Material: Scientific Inquiry When. H. Morgan s group ( he Fly Room ) showed that genes are located on chromosomes, the two components of chromosomes DN and protein became candidates for the genetic material he role of DN in heredity was first discovered by studying bacteria and the viruses that infect them

2 00 nm Evidence hat DN an ransform Bacteria he discovery of the genetic role of DN began with research by Frederick riffith in 98 Experiment Living S cells (pathogenic control) Living R cells (nonpathogenic control) Heat-killed S cells (nonpathogenic control) Mixture of heatkilled S cells and living R cells riffith worked with two strains of a bacterium (Streptococcus pneumonia), one pathogenic and one harmless When he mixed heat-killed remains of the pathogenic strain with living cells of the harmless strain, some living cells became pathogenic He called this phenomenon transformation, now defined as a change in genotype and phenotype due to assimilation of foreign DN Results Mouse dies Mouse healthy Mouse healthy Mouse dies Figure 6. an a genetic trait be transferred between different bacterial strains (riffith s experiment)? Living S cells very, Mcarty, & MacLeod In 944, Oswald very, Maclyn Mcarty, and olin MacLeod announced that the transforming substance was DN! Many biologists remained skeptical, mainly because little was known about DN Evidence hat Viral DN an Program ells More evidence for DN as the genetic material came from studies of viruses that infect bacteria Such viruses, called bacteriophages (or phages), are widely used in molecular genetics research virus is DN (sometimes RN) enclosed by a protective coat, often simply protein Figure 6.3 Virus infecting a bacterial cell Phage head Bacteriophages infecting a bacterial cell Phage head DN ail sheath ail fiber enetic material Bacterial cell Bacterial cell ail sheath ail fiber DN

3 nimation: Phage Reproductive ycle Hershey & hase: Is protein or DN the genetic material of phage? In 95, lfred Hershey and Martha hase showed that DN is the genetic material of a phage known as hey designed an experiment showing that only one of the two components of (DN or protein) enters an E. coli cell during infection hey concluded that the injected DN of the phage provides the genetic information Figure 6.4 Is protein or DN the genetic material of phage? Experiment Batch : Radioactive sulfur ( 35 S) in phage protein Labeled phages gitation frees outside infect cells. phage parts from cells. Radioactive protein 3 entrifuged cells form a pellet. 4 Radioactivity (phage protein) found in liquid nimation: Hershey-hase Experiment entrifuge Batch : Radioactive phosphorus ( 3 P) in phage DN Radioactive DN Pellet entrifuge Pellet 4 Radioactivity (phage DN) found in pellet dditional Evidence hat DN Is the enetic Material It was known that DN is a polymer of nucleotides, each consisting of a nitrogenous base, a sugar, and a phosphate group In 950, Erwin hargaff reported that DN composition varies from one species to the next his evidence of diversity made DN a more credible candidate for the genetic material hargaff s Rules wo findings became known as hargaff s Rules he base composition of DN varies between species In any species the number of and bases are equal and the number of and bases are equal he basis for these rules was not understood until the discovery of the double helix 3

4 Sugar phosphate backbone end Nitrogenous bases hymine () nimation: DN and RN Structure Figure 6.5 he structure of a DN denine () ytosine () Phosphate end Sugar DN (deoxyribose) nucleotide Nitrogenous base uanine () Building a Structural Model of DN: Scientific Inquiry fter DN was accepted as the genetic material, the challenge was to determine how its structure accounts for its role in heredity Figure 6.6 Rosalind Franklin and her X-ray diffraction photo of DN Maurice Wilkins and Rosalind Franklin were using a technique called X-ray crystallography to study molecular structure Franklin produced a picture of the DN molecule using this technique (a) Rosalind Franklin (b) Franklin s X-ray diffraction photograph of DN Building a Structural Model of DN, cont. nimation: DN Double Helix Franklin s X-ray crystallographic images of DN enabled Watson to deduce that DN was helical he X-ray images also enabled Watson to deduce the width of the helix and the spacing of the nitrogenous bases he pattern in the photo suggested that the DN molecule was made up of two s, forming a double helix 4

5 Figure 6.7 he structure of the double helix end Hydrogen bond end 3.4 nm Video: Stick Model of DN (Deoxyribonucleic cid) nm (a) Key features of DN structure 0.34 nm end (b) Partial chemical structure end (c) Space-filling model Building a Structural Model of DN, cont. Watson and rick built models of a double helix to conform to the X-rays and chemistry of DN Franklin had concluded that there were two outer sugar-phosphate backbones, with the nitrogenous bases paired in the molecule s interior Watson built a model in which the backbones were antiparallel (their subunits run in opposite directions) Building a Structural Model of DN, cont. t first, Watson and rick thought the bases paired like with like ( with, and so on), but such pairings did not result in a uniform width Instead, pairing a purine with a pyrimidine resulted in a uniform width consistent with the X-ray data Purine + purine: too wide Pyrimidine + pyrimidine: too narrow Purine + pyrimidine: width consistent with X-ray data Building a Structural Model of DN, cont. Watson and rick reasoned that the pairing was more specific, dictated by the base structures hey determined that adenine () paired only with thymine (), and guanine () paired only with cytosine () he Watson-rick model explains hargaff s rules: in any organism the amount of =, and the amount of = Figure 6.8 Base pairing in DN Sugar Sugar denine () Sugar hymine () Sugar uanine () ytosine () 5

6 oncept 6.: Many proteins work together in DN replication and repair he relationship between structure and function is manifest in the double helix Watson and rick noted that the specific base pairing suggested a possible copying mechanism for genetic material We now call this copying mechanism DN replication he Basic Principle: Base Pairing to a emplate Strand Since the two s of DN are complementary, each acts as a template for building a new in replication In DN replication, the parent molecule unwinds, and two new daughter s are built based on basepairing rules Watson and rick s semiconservative model of replication predicts that when a double helix replicates, each daughter molecule will have one old (derived or conserved from the parent molecule) and one newly made 04 Pearson Education, Inc. nimation: DN Replication Overview Figure 6.9 model for DN replication: the basic concept (a) Parental molecule (b) Separation of parental s into templates (c) Formation of new s complementary to template s DN Replication: loser Look he copying of DN is remarkable in its speed and accuracy More than a dozen enzymes and other proteins participate in DN replication etting Started Replication begins at particular sites called origins of replication, where the two DN s are separated, opening up a replication bubble eukaryotic chromosome may have hundreds or even thousands of origins of replication Replication proceeds in both directions from each origin, until the entire molecule is copied 6

7 0.5 µm 0.5 µm Figure 6. (a) Origin of replication in an E. coli cell Origin of Parental (template) replication Daughter (new) Replication Bacterial fork chromosome Doubleed Replication bubble DN molecule (b) Origins of replication in a eukaryotic cell Origin of replication Double-ed DN molecule Bubble Eukaryotic chromosome Parental (template) Daughter (new) Replication fork nimation: Origins of Replication wo daughter DN molecules wo daughter DN molecules etting Started, cont. t the end of each replication bubble is a replication fork, a Y-shaped region where new DN s are elongating Helicases are enzymes that untwist the double helix at the replication forks Single- binding proteins bind to and stabilize single-ed DN opoisomerase corrects overwinding ahead of replication forks by breaking, swiveling, and rejoining DN s Figure 6.3 Some of the proteins involved in the initiation of DN replication Primase opoisomerase Helicase Replication fork Single- binding proteins RN primer etting Started, cont. DN polymerases cannot initiate synthesis of a polynucleotide; they can only add nucleotides to an existing end he initial nucleotide is a short RN primer n enzyme called primase can start an RN chain from scratch and adds RN nucleotides one at a time using the parental DN as a template he primer is short (5 0 nucleotides long), and the end serves as the starting point for the new DN Synthesizing a New DN Strand Enzymes called DN polymerases catalyze the elongation of new DN at a replication fork Most DN polymerases require a primer and a DN template he rate of elongation is about 500 nucleotides per second in bacteria and 50 per second in human cells 7

8 New emplate ntiparallel Elongation Leading Strand Sugar Phosphate DN polymerase OH P P i Pyrophosphate Nucleotide Figure 6.4 Incorporation of a nucleotide into a DN P i OH Base he antiparallel structure of the double helix affects replication DN polymerases add nucleotides only to the free end of a growing ; therefore, a new DN can elongate only in the to direction long one template of DN, the DN polymerase synthesizes a leading continuously, moving toward the replication fork nimation: Leading Strand Figure 6.5 Synthesis of the leading during DN replication Leading Overview Origin of replication Primer Overall directions of replication Leading Figure 6.5 Synthesis of the leading during DN replication DN pol III starts to synthesize leading. Parental DN Origin of replication RN primer Sliding clamp DN pol III ntiparallel Elongation Strand o elongate the other new, called the lagging, DN polymerase must work in the direction away from the replication fork he lagging is synthesized as a series of segments called Okazaki fragments, which are later joined together by DN ligase ontinuous elongation in the to direction 8

9 Figure 6.5 Synthesis of the lagging Leading Overview Origin of replication Figure 6.5 Synthesis of the lagging Leading Overview Origin of replication Overall directions of replication Leading emplate 3 Primase makes RN primer. Origin of replication RN primer for fragment DN pol III detaches. Overall directions of replication Okazaki fragment Leading DN pol III makes Okazaki fragment. RN primer for fragment Okazaki fragment DN pol III makes Okazaki fragment. DN pol I replaces RN with DN. DN ligase forms bonds between DN fragments. Overall direction of replication nimation: Strand Figure 6.7 Summary of bacterial DN replication. Single- binding proteins Helicase Parental DN Leading template Leading DN pol III Primer Primase 5 DN pol III 4 Leading Overview Origin of replication Overall directions of replication 3 Leading DN pol I DN ligase template nimation: DN Replication Review 9

10 he DN Replication omplex he proteins that participate in DN replication form a large complex, a DN replication machine he DN replication machine may be stationary during the replication process Recent studies support a model in which DN polymerase molecules reel in parental DN and extrude newly made daughter DN molecules Figure 6.8 current model of the DN replication complex onnecting protein DN pol III template Parental DN Leading template DN pol III Helicase Leading BioFlix: DN Replication Proofreading and Repairing DN DN polymerases proofread newly made DN, replacing any incorrect nucleotides In mismatch repair of DN, repair enzymes correct errors in base pairing DN can be damaged by exposure to harmful chemical or physical agents such as cigarette smoke and X-rays; it can also undergo spontaneous changes In nucleotide excision repair, a nuclease cuts out and replaces damaged stretches of DN Figure 6.9 Nucleotide excision repair of DN damage Nuclease DN polymerase DN ligase Evolutionary Significance of ltered DN Nucleotides Error rate after proofreading repair is low, but not zero Sequence changes may become permanent and can be passed on to the next generation hese changes (mutations) are the source of the genetic variation upon which natural selection operates 0

11 Replicating the Ends of DN Molecules he limitations of DN polymerase create problems for the linear DN of eukaryotic chromosomes he usual replication machinery provides no way to complete the ends, so repeated rounds of replication produce shorter DN molecules with uneven ends his is not a problem for prokaryotes, most of which have circular chromosomes. Figure 6.0 Shortening of the ends of linear DN molecules Ends of parental DN s Parental Last fragment RN primer New leading New lagging Leading Next-to-last fragment Removal of primers and replacement with DN where a end is available Second round of replication Further rounds of replication Shorter and shorter daughter molecules elomeres Figure 6. elomeres in Mouse hromosomes Eukaryotic chromosomal DN molecules have special nucleotide sequences at their ends called telomeres elomeres do not prevent the shortening of DN molecules, but they do postpone the erosion of genes near the ends of DN molecules It has been proposed that the shortening of telomeres is connected to aging µm elomerase If chromosomes of germ cells became shorter in every cell cycle, essential genes would eventually be missing from the gametes they produce! n enzyme called telomerase catalyzes the lengthening of telomeres in germ cells he shortening of telomeres might protect cells from cancerous growth by limiting the number of cell divisions here is evidence of telomerase activity in cancer cells, which may allow cancer cells to persist oncept 6.3: chromosome consists of a DN molecule packed together with proteins he bacterial chromosome is a double-ed, circular DN molecule associated with a small amount of protein In a bacterium, the DN is supercoiled and found in a region of the cell called the nucleoid Eukaryotic chromosomes have linear DN molecules associated with a large amount of protein

12 hromatin Figure 6. hromatin packing in a eukaryotic chromosome In the eukaryotic cell, DN is precisely combined with proteins in a complex called chromatin hromosomes fit into the nucleus through an elaborate, multilevel system of packing Histones are proteins that are responsible for the first level of DN packing in chromatin. DN double helix ( nm in diameter) Histones DN, the Histones double helix Nucleosome (0 nm in diameter) Histone tail H Nucleosomes, or beads on a string (0-nm fiber) 30-nm fiber 30-nm fiber Loops Scaffold 300-nm fiber hromatid (700 nm) Replicated Looped chromosome domains (,400 nm) (300-nm fiber) Metaphase chromosome nimation: DN Packing hromatin, cont. hromatin undergoes changes in packing during the cell cycle t interphase, some chromatin is organized into a 0-nm fiber, but much is compacted into a 30-nm fiber, through folding and looping Interphase chromosomes occupy specific restricted regions in the nucleus and the fibers of different chromosomes do not become entangled Figure 6.3 Painting hromosomes hromatin, cont. Most chromatin is loosely packed in the nucleus during interphase and condenses prior to mitosis Loosely packed chromatin is called euchromatin During interphase a few regions of chromatin (centromeres and telomeres) are highly condensed into heterochromatin Dense packing of the heterochromatin makes it difficult for the cell to express genetic information coded in these regions 5 µm

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