Essential Cell Biology 5th edition
A:18 Answersuse of the word dogma, which is a belief that cannot bedoubted. I did appreciate this in a vague sort of way butsince I thought that all religious beliefs were without seriousfoundation, I used the word in the way I myself thoughtabout it, not as the world does, and simply applied it to agrand hypothesis that, however plausible, had little directexperimental support at the time.” (Francis Crick, What MadPursuit: A Personal View of Scientific Discovery. Basic Books,1988.)ANSWER 7–2 Actually, the RNA polymerases are notmoving at all in the micrograph, because they have beenfixed and coated with metal to prepare the sample forviewing in the electron microscope. However, before theywere fixed, they were moving from left to right, as indicatedby the gradual lengthening of the RNA transcripts. The RNAtranscripts are not fully extended because they begin tofold up and interact with proteins as they are synthesized;this is why they are shorter than the corresponding DNAsegments.ANSWER 7–3 At first glance, the catalytic activities ofan RNA polymerase used for transcription could replacethe primase that operates during DNA replication. Uponfurther reflection, however, there would be some seriousproblems. (1) The RNA polymerase used to make primerswould need to initiate every few hundred bases, which ismuch more often than promoters are spaced on the DNA.Initiation would therefore need to occur in a promoterindependentfashion or many more promoters wouldhave to be present in the DNA, both of which would beproblematic for the synthesis of mRNA. In addition, RNApolymerase normally begins transcription on doublestrandedDNA, whereas the DNA replication primers aresynthesized using single-stranded DNA. (2) Similarly, theRNA primers used in DNA replication are much shorter thanmRNAs. The RNA polymerase would therefore need toterminate much more frequently than during transcription.Termination would need to occur spontaneously (i.e.,without requiring a terminator sequence in the DNA) or elsemany more terminators would need to be present. Again,both of these scenarios would be problematic for mRNAproduction. Although it might be possible to overcome thisproblem if special control proteins became attached to RNApolymerase during replication, the problem has been solvedby the evolution of separate enzymes with specializedproperties. Some small DNA viruses, however, do utilizethe host RNA polymerase to make RNA primers for theirreplication.ANSWER 7–4 This experiment demonstrates that, oncean amino acid has been coupled to a tRNA, the ribosomewill trust the tRNA and “blindly” incorporate that aminoacid into the position according to the match betweenthe codon and anticodon. We can therefore conclude thata significant part of the correct reading of the geneticcode—that is, the matching of a codon in an mRNA withthe correct amino acid—is performed by the synthetaseenzymes that correctly match tRNAs and amino acids.ANSWER 7–5 The mRNA will have a 5ʹ-to-3ʹ polarity,opposite to that of the DNA strand that serves asthe template. Thus the mRNA sequence will read5ʹ-GAAAAAAGCCGUUAA-3ʹ. The N-terminal amino acidcoded for by GAA is glutamic acid. UAA specifies a stopcodon, so the C-terminal amino acid is coded for by CGUand is an arginine. Note that the usual convention indescribing the sequence of a gene is to give the sequenceof the DNA strand that is not used as a template for RNAsynthesis; this sequence is the same as that of the RNAtranscript, with T written in place of U.ANSWER 7–6 The first statement is probably correct:RNA is thought to have been the first self-replicatingcatalyst and, in modern cells, is no longer self-replicating.We can debate, however, whether this represents a “loss.”RNA now serves many roles in the cell: as messengers,as adaptors for protein synthesis, as primers for DNAreplication, as regulators of gene expression, and ascatalysts for some of the most important reactions,including RNA splicing and protein synthesis.ANSWER 7–7A. False. Ribosomes can make any protein that is specifiedby the particular mRNA that they are translating. Aftertranslation, ribosomes are released from the mRNA andcan then start translating a different mRNA. It is true,however, that a ribosome can only make one type ofprotein at a time.B. False. mRNAs are translated as linear polymers; thereis no requirement that they have any particular foldedstructure. In fact, such structures that are formed bymRNA can inhibit its translation, because the ribosomehas to unfold the mRNA in order to read the message itcontains.C. False. Ribosomal subunits can exchange partners aftereach round of translation. After a ribosome is releasedfrom an mRNA, its two subunits dissociate and enter apool of free small and large subunits from which newribosomes assemble around a new mRNA.D. False. Ribosomes are not individually enclosed in amembrane.E. False. The position of the promoter determines thedirection in which transcription proceeds and thereforewhich of the two DNA strands is used as the template.Transcription of the other strand would produce anmRNA with a completely different (and in most casesmeaningless) sequence.F. False. RNA contains uracil but not thymine.G. False. The level of a protein depends on its rate ofsynthesis and degradation but not on its catalyticactivity.ANSWER 7–8 Because the deletion in the LacheinmalmRNA is internal, it likely arose from incorrect splicingof the pre-mRNA. The simplest interpretation is that theLacheinmal gene contains a 173-nucleotide-long exon(labeled “E2” in Figure A7−8), and that this exon is lost(“skipped”) during the processing of the mutant precursormRNA (pre-mRNA). This could occur, for example, if themutation changed the 3ʹ splice site in the preceding intron(“I1”) so that it was no longer recognized by the splicingmachinery (a change in the CAG sequence shown in Figure7–20 could do this). The snRNP would search for the nextavailable 3ʹ splice site, which is found at the 3ʹ end of thenext intron (“I2”), and the splicing reaction would thereforeremove E2 together with I1 and I2, resulting in a shortenedmRNA. The mRNA is then translated into a defectiveprotein, resulting in the Lacheinmal deficiency.Because 173 nucleotides do not amount to an integral
Answers A:19(A) NORMALsplicing173 bpsplicing5′ 3′cap E1 E2 E3 AAApre-mRNA(B) MUTANTFigure A7−8E1 I1 E2 I2 E3capE3 AAAmRNALacheinmal proteingenemutation that inactivates 3′ splice siteE1 I1 E2 I2 E3mutant genesplicing5′ 3′cap E1 E2 E3 AAAmutant pre-mRNAcapE1E1E2mutant proteinE3 AAAmutant RNAnumber of codons, the lack of this exon in the mRNA willshift the reading frame at the splice junction. Therefore, theLacheinmal protein would be made correctly only throughexon E1. As the ribosome begins translating sequencesin exon E3, it will be in the wrong reading frame and willtherefore will produce a protein sequence that is unrelatedto the Lacheinmal sequence normally encoded by exon E3.ECB5 EA7.08/Most likely, the ribosome will soon encounter a stop codon,which would be expected to occur on average about once inevery 21 codons (there are 3 stop codons in the 64 codonsof the genetic code).ANSWER 7–9 Sequence 1 and sequence 4 both codefor the peptide Arg-Gly-Asp. Because the genetic code isredundant, different nucleotide sequences can encode thesame amino acid sequence.ANSWER 7–10A. Incorrect. The bonds are not covalent, and theirformation does not require an input of energy.B. Correct. The aminoacyl-tRNA enters the ribosome at theA site and forms hydrogen bonds with the codon in themRNA.C. Correct. As the ribosome moves along the mRNA,the tRNAs that have donated their amino acid tothe growing polypeptide chain are ejected from theribosome and the mRNA. The ejection takes place twocycles after the tRNA first enters the ribosome (seeFigure 7–37).ANSWER 7–11 Replication. Dictionary definition: thecreation of an exact copy; molecular biology definition: theact of copying a DNA sequence. Transcription. Dictionarydefinition: the act of writing out a copy, especially from onephysical form to another; molecular biology definition: theact of copying the information stored in DNA into RNA.Translation. Dictionary definition: the act of putting wordsinto a different language; molecular biology definition:the act of polymerizing amino acids into a defined linearsequence using the information provided by the linearsequence of nucleotides in mRNA. (Note that “translation”is also used in a quite different sense, both in ordinarylanguage and in scientific contexts, to mean a movementfrom one place to another.)ANSWER 7–12 With four different nucleotides to choosefrom, a code of two nucleotides could specify 16 differentamino acids (= 4 2 ), and a triplet code in which the positionof the nucleotides is not important could specify 20 differentamino acids (= 4 possibilities of 3 of the same bases +12 possibilities of 2 bases the same and one different +4 possibilities of 3 different bases). In both cases, thesemaximal amino acid numbers would need to be reduced byat least 1 because of the need to specify translation stopcodons. It is relatively easy to envision how a doublet codecould be translated by a mechanism similar to that used inour world by providing tRNAs with only two relevant basesin the anticodon loop. It is more difficult to envision howthe nucleotide composition of a stretch of three nucleotidescould be translated without regard to their order, becausebase-pairing can then no longer be used: AUG, for example,will not base-pair with the same anticodon as UGA.ANSWER 7–13 It is likely that in early cells the matchingbetween codons and amino acids was less accurate thanit is in present-day cells. The feature of the genetic codedescribed in the question may have allowed early cells totolerate this inaccuracy by allowing a blurred relationshipbetween sets of roughly similar codons and roughly similaramino acids. One can easily imagine how the matchingbetween codons and amino acids could have become moreaccurate, step by step, as the translation machinery evolvedinto that found in modern cells.ANSWER 7–14 The codon for Trp is 5ʹ-UGG-3ʹ. Thusa normal tRNA Trp contains the sequence 5ʹ-CCA-3ʹ asits anticodon (see Figure 7–33). If this tRNA contains amutation so that its anticodon is changed to UCA, it willrecognize a UGA codon and lead to the incorporation ofa tryptophan instead of causing translation to stop. Manyother protein-encoding sequences, however, contain UGAcodons as their natural stop sites, and these stops wouldalso be affected by the mutant tRNA. Depending on thecompetition between the altered tRNA and the normaltranslation release factors (Figure 7–41), some of theseproteins would be made with additional amino acids at theirC-terminal end. The additional lengths would depend on thenumber of codons before the ribosomes encounter a non-UGA stop codon in the mRNA in the reading frame in whichthe protein is translated.ANSWER 7–15 One effective way of driving a reaction tocompletion is to remove one of the products, so that thereverse reaction cannot occur. ATP contains two high-energybonds that link the three phosphate groups. In the reactionshown, PP i is released, consisting of two phosphate groupslinked by one of these high-energy bonds. Thus PP i can behydrolyzed with a considerable gain of free energy, andthereby can be efficiently removed. This happens rapidly in
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A:18 Answers
use of the word dogma, which is a belief that cannot be
doubted. I did appreciate this in a vague sort of way but
since I thought that all religious beliefs were without serious
foundation, I used the word in the way I myself thought
about it, not as the world does, and simply applied it to a
grand hypothesis that, however plausible, had little direct
experimental support at the time.” (Francis Crick, What Mad
Pursuit: A Personal View of Scientific Discovery. Basic Books,
1988.)
ANSWER 7–2 Actually, the RNA polymerases are not
moving at all in the micrograph, because they have been
fixed and coated with metal to prepare the sample for
viewing in the electron microscope. However, before they
were fixed, they were moving from left to right, as indicated
by the gradual lengthening of the RNA transcripts. The RNA
transcripts are not fully extended because they begin to
fold up and interact with proteins as they are synthesized;
this is why they are shorter than the corresponding DNA
segments.
ANSWER 7–3 At first glance, the catalytic activities of
an RNA polymerase used for transcription could replace
the primase that operates during DNA replication. Upon
further reflection, however, there would be some serious
problems. (1) The RNA polymerase used to make primers
would need to initiate every few hundred bases, which is
much more often than promoters are spaced on the DNA.
Initiation would therefore need to occur in a promoterindependent
fashion or many more promoters would
have to be present in the DNA, both of which would be
problematic for the synthesis of mRNA. In addition, RNA
polymerase normally begins transcription on doublestranded
DNA, whereas the DNA replication primers are
synthesized using single-stranded DNA. (2) Similarly, the
RNA primers used in DNA replication are much shorter than
mRNAs. The RNA polymerase would therefore need to
terminate much more frequently than during transcription.
Termination would need to occur spontaneously (i.e.,
without requiring a terminator sequence in the DNA) or else
many more terminators would need to be present. Again,
both of these scenarios would be problematic for mRNA
production. Although it might be possible to overcome this
problem if special control proteins became attached to RNA
polymerase during replication, the problem has been solved
by the evolution of separate enzymes with specialized
properties. Some small DNA viruses, however, do utilize
the host RNA polymerase to make RNA primers for their
replication.
ANSWER 7–4 This experiment demonstrates that, once
an amino acid has been coupled to a tRNA, the ribosome
will trust the tRNA and “blindly” incorporate that amino
acid into the position according to the match between
the codon and anticodon. We can therefore conclude that
a significant part of the correct reading of the genetic
code—that is, the matching of a codon in an mRNA with
the correct amino acid—is performed by the synthetase
enzymes that correctly match tRNAs and amino acids.
ANSWER 7–5 The mRNA will have a 5ʹ-to-3ʹ polarity,
opposite to that of the DNA strand that serves as
the template. Thus the mRNA sequence will read
5ʹ-GAAAAAAGCCGUUAA-3ʹ. The N-terminal amino acid
coded for by GAA is glutamic acid. UAA specifies a stop
codon, so the C-terminal amino acid is coded for by CGU
and is an arginine. Note that the usual convention in
describing the sequence of a gene is to give the sequence
of the DNA strand that is not used as a template for RNA
synthesis; this sequence is the same as that of the RNA
transcript, with T written in place of U.
ANSWER 7–6 The first statement is probably correct:
RNA is thought to have been the first self-replicating
catalyst and, in modern cells, is no longer self-replicating.
We can debate, however, whether this represents a “loss.”
RNA now serves many roles in the cell: as messengers,
as adaptors for protein synthesis, as primers for DNA
replication, as regulators of gene expression, and as
catalysts for some of the most important reactions,
including RNA splicing and protein synthesis.
ANSWER 7–7
A. False. Ribosomes can make any protein that is specified
by the particular mRNA that they are translating. After
translation, ribosomes are released from the mRNA and
can then start translating a different mRNA. It is true,
however, that a ribosome can only make one type of
protein at a time.
B. False. mRNAs are translated as linear polymers; there
is no requirement that they have any particular folded
structure. In fact, such structures that are formed by
mRNA can inhibit its translation, because the ribosome
has to unfold the mRNA in order to read the message it
contains.
C. False. Ribosomal subunits can exchange partners after
each round of translation. After a ribosome is released
from an mRNA, its two subunits dissociate and enter a
pool of free small and large subunits from which new
ribosomes assemble around a new mRNA.
D. False. Ribosomes are not individually enclosed in a
membrane.
E. False. The position of the promoter determines the
direction in which transcription proceeds and therefore
which of the two DNA strands is used as the template.
Transcription of the other strand would produce an
mRNA with a completely different (and in most cases
meaningless) sequence.
F. False. RNA contains uracil but not thymine.
G. False. The level of a protein depends on its rate of
synthesis and degradation but not on its catalytic
activity.
ANSWER 7–8 Because the deletion in the Lacheinmal
mRNA is internal, it likely arose from incorrect splicing
of the pre-mRNA. The simplest interpretation is that the
Lacheinmal gene contains a 173-nucleotide-long exon
(labeled “E2” in Figure A7−8), and that this exon is lost
(“skipped”) during the processing of the mutant precursor
mRNA (pre-mRNA). This could occur, for example, if the
mutation changed the 3ʹ splice site in the preceding intron
(“I1”) so that it was no longer recognized by the splicing
machinery (a change in the CAG sequence shown in Figure
7–20 could do this). The snRNP would search for the next
available 3ʹ splice site, which is found at the 3ʹ end of the
next intron (“I2”), and the splicing reaction would therefore
remove E2 together with I1 and I2, resulting in a shortened
mRNA. The mRNA is then translated into a defective
protein, resulting in the Lacheinmal deficiency.
Because 173 nucleotides do not amount to an integral