Essential Cell Biology 5th edition
A:14 Answersthe sugar–phosphate backbone is vastly different in theA-C pair compared with A-T, and the spacing between thetwo sugar–phosphate strands is considerably increasedin the A-G pair, where two large purine rings interact.Consequently, it is energetically unfavorable to incorporatea wrong base in DNA, and such errors occur only very rarely.ANSWER 5–7A. The bases V, W, X, and Y can form a DNA-like doublehelicalmolecule with virtually identical properties tothose of bona fide DNA. V would always pair with X,and W with Y. Therefore, the macromolecule couldbe derived from a living organism that uses the sameprinciples to replicate its genome as those used byorganisms on Earth. In principle, different bases, suchas V, W, X, and Y, could have been selected duringevolution on Earth as building blocks for DNA. (Similarly,there are many more conceivable amino acid side chainsthan the set of 20 selected in evolution that make up allproteins.)B. None of the bases V, W, X, or Y can replace A, T, G, orC. To preserve the distance between the two sugar–phosphate strands in a double helix, a pyrimidine alwayshas to pair with a purine (see, for example, Figure 5–4).Thus, the eight possible combinations would be V-A,V-G, W-A, W-G, X-C, X-T, Y-C, and Y-T. Because of thepositions of hydrogen-bond acceptors and hydrogenbonddonor groups, however, no stable base pairs wouldform in any of these combinations, as shown for thepairing of V and A in Figure A5–7, where only a singlehydrogen bond could form.Figure A5−7HVNHHadenineNCCNHCNNCNANSWER 5–8 As the two strands are held togetherby hydrogen bonds between the bases, the stability of aDNA double helix is largely dependent on the number ofhydrogen bonds that can be formed. Thus two parametersdetermine the stability: the number of nucleotide pairs andthe number of hydrogen bonds that each nucleotide paircontributes. As shown in Figure 5–4, an A-T pair contributestwo hydrogen bonds, whereas a G-C pair contributes threehydrogen bonds. Therefore, helix C (containing a total of34 hydrogen bonds) would melt at the lowest temperature,helix B (containing a total of 65 hydrogen bonds) would meltCCCCCHOHNNHHnext, and helix A (containing a total of 78 hydrogen bonds)would melt last. Helix A is the most stable, largely owingto its high GC content. Indeed, the DNA of organisms thatgrow in extreme temperature environments, such as certainprokaryotes that grow in geothermal vents, has an unusuallyhigh GC content.ANSWER 5–9 The DNA would be enlarged by a factorof 2.5 × 10 6 (= 5 × 10 –3 /2 × 10 –9 m). Thus the extensioncord would be 2500 km long. This is approximately thedistance from London to Istanbul, San Francisco to KansasCity, Tokyo to the southern tip of Taiwan, and Melbourne toCairns. Adjacent nucleotides would be about 0.85 mm apart(which is only about the thickness of a stack of 12 pages ofthis book). A gene that is 1000 nucleotide pairs long wouldbe about 85 cm in length.ANSWER 5–10A. It takes two bits to specify each nucleotide pair (forexample, 00, 01, 10, and 11 would be the binary codesfor the four different nucleotides, each paired with itsappropriate partner).B. The entire human genome (3 × 10 9 nucleotide pairs)could be stored on two CDs (3 × 10 9 × 2 bits/4.8 × 10 9bits).ANSWER 5–11A. True.B. False. Nucleosome core particles are approximately11 nm in diameter.ANSWER 5–12 The definitions of the terms can be foundin the Glossary. DNA assembles with specialized proteinsto form chromatin. At a first level of packing, histones formthe core of nucleosomes. A nucleosome includes the DNAwrapped around this histone core plus a segment of linkerDNA. Between nuclear divisions—that is, in interphase—thechromatin of the interphase chromosomes is in a relativelyextended form in the nucleus, although some regions ofit, the heterochromatin, remain densely packed and aretranscriptionally inactive. During nuclear division—that is, inmitosis—replicated chromosomes become condensed intomitotic chromosomes, which are transcriptionally inactiveand are designed to be readily distributed between the twodaughter cells.ANSWER 5–13 Colonies are clumps of cells that originatefrom a single founder cell and grow outward as the cellsdivide again and again. In the lower colony of Figure Q5–13,the Ade2 gene is inactivated when placed near a telomere,but apparently it can become spontaneously activated in afew cells, which then turn white. Once activated in a cell, theAde2 gene continues to be active in the descendants of thatcell, resulting in clumps of white cells (the white sectors)in the colony. This result shows both that the inactivationof a gene positioned close to a telomere can be reversedand that this change is passed on to further generations.This change in Ade2 expression probably results from aspontaneous decondensation of the chromatin structurearound the gene.ANSWER 5–14 In the electron micrographs, one candetect chromatin regions of two different densities; thedensely stained regions correspond to heterochromatin,while less condensed chromatin is more lightly stained.The chromatin in (A) is mostly in the form of condensed,
Answers A:15transcriptionally inactive heterochromatin, whereas mostof the chromatin in (B) is decondensed and thereforepotentially transcriptionally active. The nucleus in (A) isfrom a reticulocyte, a red blood cell precursor, which islargely devoted to making a single protein, hemoglobin.The nucleus in (B) is from a lymphocyte, which is active intranscribing many different genes.ANSWER 5–15 Helix (A) is right-handed. Helix (C) isleft-handed. Helix (B) has one right-handed strand andone left-handed strand. There are several ways to tell thehandedness of a helix. For a vertically oriented helix, likethe ones in Figure Q5–15, if the strands in front point upto the right, the helix is right-handed; if they point up tothe left, the helix is left-handed. Once you are comfortableidentifying the handedness of a helix, you will be amusedto note that nearly 50% of the “DNA” helices shownin advertisements are left-handed, as are a surprisinglyhigh number of the ones shown in books. Amazingly,a version of Helix (B) was used in advertisements for aprominent international conference, celebrating the 30-yearanniversary of the discovery of the DNA helix.ANSWER 5–16 The packing ratio within a nucleosome coreis 4.5 [(147 bp × 0.34 nm/bp)/(11 nm) = 4.5]. If there is anadditional 54 bp of linker DNA, then the packing ratio for“beads-on-a-string” DNA is 2.3 [(201 bp × 0.34 nm/bp)/(11 nm + {54 bp × 0.34 nm/bp}) = 2.3]. This first level ofpacking represents only 0.023% (2.3/10,000) of the totalcondensation that occurs at mitosis.Chapter 6ANSWER 6–1A. The distance between replication forks 4 and 5 is about280 nm, corresponding to 824 nucleotides (= 280/0.34).These two replication forks would collide in about 8seconds. Forks 7 and 8 move away from each other andwould therefore never collide.B. The total length of DNA shown in the electronmicrograph is about 1.5 μm, corresponding to 4400nucleotides. This is only about 0.002%[= (4400/1.8 × 10 8 ) × 100%] of the total DNA in a fly cell.ANSWER 6–2 Although the process may seem wasteful,it is not possible to proofread during the initial stages ofprimer synthesis. To start a new primer on a piece of singlestrandedDNA, one nucleotide needs to be put in placeand then linked to a second, and then to a third, and so on.Even if these first nucleotides were perfectly matched tothe template strand, they would bind with very low affinity,and it would consequently be difficult for a hypotheticalprimase with proofreading activity to distinguish the correctfrom incorrect bases; the enzyme would therefore stall.The task of the primase is to “just polymerize nucleotidesthat bind reasonably well to the template without worryingtoo much about accuracy.” Later, these sequences areremoved and replaced by DNA polymerase, which usesnewly synthesized, adjacent DNA—which has already beenproofread—as its primer.ANSWER 6–3A. Without DNA polymerase, no replication can take placeat all. RNA primers will be laid down at the origin ofreplication.B. DNA ligase links the DNA fragments that are producedon the lagging strand. In the absence of ligase, thenewly replicated DNA strands will remain as fragments,but no nucleotides will be missing.C. Without the sliding clamp, the DNA polymerase willfrequently fall off the DNA template. In principle, it canrebind and continue, but the continual falling off andrebinding will be so time-consuming that the cell will beunable to divide.D. In the absence of RNA-excision enzymes, the RNAfragments will remain covalently attached to the newlyreplicated DNA fragments. No ligation will take place,because the DNA ligase will not link DNA to RNA.The lagging strand will therefore consist of fragmentscomposed of both RNA and DNA.E. Without DNA helicase, the DNA polymerase willstall because it cannot separate the strands of thetemplate DNA ahead of it. Little or no new DNA will besynthesized.F. In the absence of primase, RNA primers cannot bemade on either the leading or the lagging strand. DNAreplication therefore cannot begin.ANSWER 6–4 DNA damage by deamination anddepurination reactions occurs spontaneously. This typeof damage is not the result of replication errors and istherefore equally likely to occur on either strand. If DNArepair enzymes recognized such damage only on newlysynthesized DNA strands, half of the defects would gouncorrected. The statement is therefore incorrect.ANSWER 6–5 If the old strand were “repaired” using thenew strand that contains a replication error as the template,then the error would become a permanent mutation inthe genome. The old information would be erased in theprocess. Therefore, if repair enzymes did not distinguishbetween the two strands, there would be only a 50% chancethat any given replication error would be corrected.ANSWER 6–6 You cannot transform an individual fromone species into another species simply by introducingrandom changes into the DNA. It is exceedingly unlikelythat the 5000 mutations that would accumulate every day inthe absence of the DNA repair enzyme would be in the verypositions where human and chimpanzee DNA sequencesare different. It is very likely that, at such a high mutationfrequency, many essential genes would be inactivated,leading to cell death. Furthermore, your body is made upof about 10 13 cells. For you to turn into an ape, not justone but many of these cells would need to be changed.And even then, many of these changes would have to occurduring development to effect changes in your body plan(making your arms longer than your legs, for example).ANSWER 6–7A. False. Identical DNA polymerase molecules catalyzeDNA synthesis on the leading and lagging strandsof a bacterial replication fork. The replication forkis asymmetrical because the lagging strand is madein pieces while the leading strand is synthesizedcontinuously.B. False. Okazaki fragments initially contain both RNAprimers and DNA, but only the RNA primers areremoved by RNA nucleases.
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A:14 Answers
the sugar–phosphate backbone is vastly different in the
A-C pair compared with A-T, and the spacing between the
two sugar–phosphate strands is considerably increased
in the A-G pair, where two large purine rings interact.
Consequently, it is energetically unfavorable to incorporate
a wrong base in DNA, and such errors occur only very rarely.
ANSWER 5–7
A. The bases V, W, X, and Y can form a DNA-like doublehelical
molecule with virtually identical properties to
those of bona fide DNA. V would always pair with X,
and W with Y. Therefore, the macromolecule could
be derived from a living organism that uses the same
principles to replicate its genome as those used by
organisms on Earth. In principle, different bases, such
as V, W, X, and Y, could have been selected during
evolution on Earth as building blocks for DNA. (Similarly,
there are many more conceivable amino acid side chains
than the set of 20 selected in evolution that make up all
proteins.)
B. None of the bases V, W, X, or Y can replace A, T, G, or
C. To preserve the distance between the two sugar–
phosphate strands in a double helix, a pyrimidine always
has to pair with a purine (see, for example, Figure 5–4).
Thus, the eight possible combinations would be V-A,
V-G, W-A, W-G, X-C, X-T, Y-C, and Y-T. Because of the
positions of hydrogen-bond acceptors and hydrogenbond
donor groups, however, no stable base pairs would
form in any of these combinations, as shown for the
pairing of V and A in Figure A5–7, where only a single
hydrogen bond could form.
Figure A5−7
H
V
N
H
H
adenine
N
C
C
N
H
C
N
N
C
N
ANSWER 5–8 As the two strands are held together
by hydrogen bonds between the bases, the stability of a
DNA double helix is largely dependent on the number of
hydrogen bonds that can be formed. Thus two parameters
determine the stability: the number of nucleotide pairs and
the number of hydrogen bonds that each nucleotide pair
contributes. As shown in Figure 5–4, an A-T pair contributes
two hydrogen bonds, whereas a G-C pair contributes three
hydrogen bonds. Therefore, helix C (containing a total of
34 hydrogen bonds) would melt at the lowest temperature,
helix B (containing a total of 65 hydrogen bonds) would melt
C
C
C
C
C
H
O
H
N
N
H
H
next, and helix A (containing a total of 78 hydrogen bonds)
would melt last. Helix A is the most stable, largely owing
to its high GC content. Indeed, the DNA of organisms that
grow in extreme temperature environments, such as certain
prokaryotes that grow in geothermal vents, has an unusually
high GC content.
ANSWER 5–9 The DNA would be enlarged by a factor
of 2.5 × 10 6 (= 5 × 10 –3 /2 × 10 –9 m). Thus the extension
cord would be 2500 km long. This is approximately the
distance from London to Istanbul, San Francisco to Kansas
City, Tokyo to the southern tip of Taiwan, and Melbourne to
Cairns. Adjacent nucleotides would be about 0.85 mm apart
(which is only about the thickness of a stack of 12 pages of
this book). A gene that is 1000 nucleotide pairs long would
be about 85 cm in length.
ANSWER 5–10
A. It takes two bits to specify each nucleotide pair (for
example, 00, 01, 10, and 11 would be the binary codes
for the four different nucleotides, each paired with its
appropriate partner).
B. The entire human genome (3 × 10 9 nucleotide pairs)
could be stored on two CDs (3 × 10 9 × 2 bits/4.8 × 10 9
bits).
ANSWER 5–11
A. True.
B. False. Nucleosome core particles are approximately
11 nm in diameter.
ANSWER 5–12 The definitions of the terms can be found
in the Glossary. DNA assembles with specialized proteins
to form chromatin. At a first level of packing, histones form
the core of nucleosomes. A nucleosome includes the DNA
wrapped around this histone core plus a segment of linker
DNA. Between nuclear divisions—that is, in interphase—the
chromatin of the interphase chromosomes is in a relatively
extended form in the nucleus, although some regions of
it, the heterochromatin, remain densely packed and are
transcriptionally inactive. During nuclear division—that is, in
mitosis—replicated chromosomes become condensed into
mitotic chromosomes, which are transcriptionally inactive
and are designed to be readily distributed between the two
daughter cells.
ANSWER 5–13 Colonies are clumps of cells that originate
from a single founder cell and grow outward as the cells
divide again and again. In the lower colony of Figure Q5–13,
the Ade2 gene is inactivated when placed near a telomere,
but apparently it can become spontaneously activated in a
few cells, which then turn white. Once activated in a cell, the
Ade2 gene continues to be active in the descendants of that
cell, resulting in clumps of white cells (the white sectors)
in the colony. This result shows both that the inactivation
of a gene positioned close to a telomere can be reversed
and that this change is passed on to further generations.
This change in Ade2 expression probably results from a
spontaneous decondensation of the chromatin structure
around the gene.
ANSWER 5–14 In the electron micrographs, one can
detect chromatin regions of two different densities; the
densely stained regions correspond to heterochromatin,
while less condensed chromatin is more lightly stained.
The chromatin in (A) is mostly in the form of condensed,