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CHAPTER 8
Manipulating Proteins, DNA, and RNA
Molecular Biology of the Cell, Fifth Edition
© 2008 Garland Science Publishing
Questions
8-1 You have accidentally torn the labels off two tubes, each containing a different
plasmid, and now do not know which plasmid is in which tube. Fortunately, you
have restriction maps for both plasmids, shown in Figure Q8-1. You have the
opportunity to test just one sample from one of your tubes. You have equipment
for agarose gel electrophoresis, a standard set of DNA size markers, and the
necessary restriction enzymes.
A. Outline briefly the series of steps you would perform to determine which
plasmid is in which tube.
B. Which restriction enzyme or combination of restriction enzymes would
you use in this experiment?
Figure Q8-1
8-2 You have sequenced a short piece of DNA and produced the gel shown below:
A. What is the sequence of the DNA, starting from the 5 end?
B. If you know that this sequence is from the middle of a protein-coding
cDNA clone, what amino acid sequence can you deduce from this
sequence?
Figure Q8-2
8-3 Figure Q8-3 shows the recognition sequences for the restriction enzymes SalI,
XhoI, PstI, and SmaI and a plasmid with the sites of cleavage for these enzymes
marked.
A. Consider all possible digestion reactions with one or two restriction
enzymes. After which of the digestions can the plasmid form into a circle
again simply by treatment with DNA ligase? Assume that after digestion
any small pieces of DNA are removed, and it is only the larger portion of
plasmid that you are trying to restore to its circular state.
B. After which of the possible digestions can the plasmid be restored to a
circle by first adding DNA polymerase and the four deoxynucleotides,
then treating with DNA ligase? Again assume that you are trying to form
the larger portion of plasmid into a circle again.
Figure Q8-3
8-4 You have an oligonucleotide probe that hybridizes to part of gene A from a
eucaryotic cell. You can use this probe to isolate a corresponding plasmid from a
DNA library harbored by a collection of bacteria. Will a cDNA library or a
genomic DNA library be more appropriate for the following applications?
Explain.
A. You want to study the promoter of a gene A.
B. Gene A encodes a tRNA and you wish to isolate a piece of DNA
containing the full-length sequence of the tRNA.
C. You discover that gene A is alternatively spliced and you want to see
which predicted alternative splice products are actually produced in a cell.
D. You want to find both gene A and the genes located near gene A on the
chromosome.
E. You want to express gene A in bacteria to produce lots of protein A.
F. You want to perform a phylogenetic comparison to find amino acid
sequences important for the function of gene A.
8-5 You want to make an antibody against a C. elegans nematode protein that the
DNA shown in Figure Q8-5A encodes. To do this, you will clone the coding
sequence into a bacterial expression plasmid, overexpress the protein, purify it,
and inject it into a mouse to stimulate the production of antibodies against the
protein.
A. You use polymerase chain reaction (PCR) amplification to amplify the
DNA corresponding to the 1800 base pairs of coding sequence; this
coding sequence lies between the two flanking sequence shown in Figure
Q8-5A. You will then insert the PCR product into the BamHI site of the
expression plasmid shown in Figure Q8-5B. Is it best to use genomic
DNA or cDNA as template in the PCR reaction? What are the sequences
of the two oligonucleotide primers that will allow you to amplify the DNA
by PCR and insert it into the BamHI site? (Remember to indicate the 5
and 3 ends of the primers.)
B. Briefly describe the series of enzymatic treatments you will use to create
the expression plasmid before transferring it into bacterial cells.
C. Once the correct new expression plasmid is in bacterial cells, you induce
high levels of expression of your nematode protein. Briefly describe the
series of steps you will perform to go from the cells containing high levels
of protein to partly purified protein.
D. Several weeks after you inject a mouse with your purified protein, you
sample its blood serum, which contains antibodies. You know from in situ
hybridizations that the mRNA corresponding to your protein is found in
gonad cells but not in gut cells. To test whether the mouse has made
antibodies against your protein, you isolate protein from gonad cells and
from gut cells and perform a Western blot (also known as an immunoblot)
with the mouse serum and a fluorescent second antibody that binds mouse
antibodies. Your result is shown in Figure Q8-5C. Did the mouse make
antibodies against your protein? Does the serum specifically recognize
only one nematode protein? (Illustrate your written answers by labeling
the blot.) What might you do next if you wanted to use antibodies from
this mouse to determine the subcellular location of your protein in fixed
cells?
Figure Q8-5
8-6 Current technologies allow much more facile manipulation and analysis of nucleic
acids than proteins. For example, it is straightforward to generate and purify
thousands of different DNA molecules for a DNA microarray, but difficult to
make an analogous protein microarray. This may reflect fundamental differences
between the two kinds of polymers. Colloquially, we might say that DNA double
helices are all the same, but each protein has its own personality.
A. Compare how the shape of a DNA double helix or protein molecule
depends on the sequence of monomers.
B. Compare the means used to detect a specific DNA molecule in a complex
mixture of other DNAs with that used to detect a specific protein in a
complex protein mixture.
C. Briefly compare the means used to make many copies of a DNA or protein
molecule for biochemical experiments.
D. Can you unambiguously predict an amino acid sequence from a cDNA
sequence? Can you unambiguously predict a cDNA sequence from an
amino acid sequence? Explain.
8-7 You are working in a laboratory, trying to identify how the nematode worm C.
elegans senses and moves toward specific “attractant” chemicals. You examine an
electronic database that compiles mRNA expression profiles of all nematode
genes measured by DNA microarray hybridization experiments, and find that an
unknown gene is transcribed coordinately with several genes known to be
involved in the attractant response. You name this gene Unk1 and set out to learn
what it does.
A. Without doing a single experiment at the lab bench, how can you learn
more about what the Unk1 protein might be doing? How might this
influence your choice of experiments with Unk1?
B. You decide to use reverse genetics to learn more about the function of the
Unk1 gene. What is the simplest first experiment to do, considering that
you are working with nematodes?
C. You decide to look for proteins that bind to Unk1, which may provide
important clues to its function. Briefly describe an approach to find
binding partners.
D. You decide to determine where and when Unk1 is expressed within the
animal, to learn more about how it causes the observed phenotypes.
Describe an approach to monitor spatiotemporal dynamics of Unk1
expression.
8-8 Human babies use the lactase enzyme to metabolize the milk sugar lactose. Most
human adults are lactose-intolerant because this gene is normally turned off in
adults. However, many adults of European descent still express lactase. This
lactase-persistence trait is tightly linked to a SNP called C/T-13910 located near
the lactase gene. Examination of many SNPs flanking the lactase gene provided
evidence that lactase persistence has been subject to very strong positive selective
pressure and became prevalent relatively recently, about 7500 years ago, which is
roughly coincident with the time that dairy farming arose in northern Europe.
A. Much of the supporting genomic evidence comes from the sizes of
haplotype blocks. Are the haplotype blocks surrounding the lactase gene
in lactase-persistent individuals bigger or smaller than the average blocks
found throughout the genome, or of average size? Are the corresponding
haplotype blocks in lactose-intolerant individuals bigger, smaller, or
average?
B. Does the C/T-13910 SNP cause the lactase-persistence trait?
C. Name an experimental technique that can determine if an unknown adult
is lactose-intolerant or lactase-persistent.
D. Is lactase persistence likely to be a dominant trait or a recessive trait?
Answers
8-1
A. First, digest DNA from one of the tubes with a combination of restriction
enzymes that will yield DNA fragments with sizes that are unique to one
of the plasmids. Then, load the resulting mixture of DNA fragments on an
agarose gel alongside a set of size markers, use electrophoresis to separate
DNAs by size, stain the gel to reveal the DNAs, and determine the
fragment sizes. By looking at the restriction maps, you should then be able
to match your results to one of the plasmids.
B. Digestion with any of the following combinations will enable you to
distinguish which plasmid you have: HindIII + BglII; EcoRI + BglII;
EcoRI + BglII + HindIII. The plasmids are the same size, so you cannot
distinguish them simply by making a single cut (with HindIII or BglII
alone) and determining the size of the complete DNA by gel
electrophoresis. Nor can you distinguish them by cutting with all four
restriction nucleases (or with BamHI and BglII), because the set of
fragment sizes produced from both plasmids will be the same. Cutting
with BamHI or EcoRI on their own is not sufficient because you will get
bands of the same size from both plasmid A and plasmid B. The only
difference between the two plasmids is the location of the BglII site
relative to the two BamHI sites, so if you cut with an enzyme that cuts
outside the BamHI fragment and with BglII, you will get different-sized
fragments from the two plasmids.
8-2
A. TAGACTGACCTG
B. Arg-Leu-Thr. You can only use the second reading frame; reading frame 1
contains a stop codon (TAG), as does reading frame 3 (TGA).
8-3
A. Ligase treatment alone can convert the large cleaved plasmid fragments to
circular forms after the plasmid is first digested with any of the single
restriction enzymes and one of the double enzyme combinations. When
SalI and XhoI cut DNA, the staggered ends left behind will match up by
base-pairing and thus can be joined by ligase alone. When all other pairs
of enzymes cut DNA, the ends will not be complementary; that is, there
will not be two sticky ends that can anneal, nor will there be two blunt
ends that can be ligated.
B. In addition to the answers for A, there are two new enzyme pairs that will
allow the plasmid to form again into a circle after treatment with DNA
polymerase and ligase: SalI + SmaI and XhoI + SmaI. Two blunt ends can
be joined by DNA ligase. Digestion with SmaI creates a blunt end.
Addition of DNA polymerase and the four deoxynucleotides will fill in the
5 overhangs generated by digestion with SalI and XhoI, leaving blunt
ends. However, 3 overhangs (namely those generated by PstI) will not be
filled in, because DNA polymerase can only add nucleotides to a recessed
3 end that is paired with a longer template strand (a 5 overhang). DNA
ligase will not join 3 overhangs from PstI digestion to any of the blunt
ends created by the other enzymes.
8-4
A. Genomic library. cDNAs are produced from mRNAs. Therefore, the
promoters will not be included in a cDNA library.
B. Genomic library. cDNAs are usually constructed by using an oligo-dT
primer, and tRNAs do not have poly-A tails. Therefore, tRNA sequences
are not usually found in cDNA libraries. If the cDNA library were made
using random primers, it would be unlikely to contain the full-length
version of the tRNA.
C. cDNA library. Because cDNAs are produced from mRNAs, isolating
cDNAs would tell you which splice variants are produced in a cell.
D. Genomic library. Some genomic DNA fragments will probably contain the
genes next to your gene of interest; cDNAs will not.
E. cDNA library. Bacteria do not have the ability to remove introns, which
may exist in DNA isolated from a genomic library.
F. cDNA library. A phylogenetic comparison of proteins requires amino acid
sequences or nucleotide coding sequences from orthologous or
homologous proteins. It is often difficult to use genomic sequences to find
unambiguously the coding sequences, sprinkled among the more abundant
intron and regulatory region sequences.
8-5
A. It is best to use cDNA as template, because the bacteria will be unable to
splice out introns in the genomic DNA. The appropriate PCR primers are
5-GGATCCGACCTGTGGAAGC-3 and
5-GGATCCTCAATCCCGTATG-3. The first primer will hybridize to
the bottom strand and prime synthesis in the rightward direction. The
second primer will hybridize to the top strand and prime synthesis in the
leftward direction. Both primers have the BamHI recognition site added to
the 5’ end to allow cleavage by BamHI, which will generate
complementary single-stranded ends that can hybridize to the ends of the
BamHI-cut plasmid.
B. First, use a thermophilic DNA polymerase for PCR amplification. Second,
use BamHI to digest both the PCR product and the plasmid. Third, use
DNA ligase to join the two digested DNA molecules together.
C. First, use ultrasonic vibration or another method to break open the cells.
Second, use preparative ultracentrifugation to remove the whole cells and
cellular debris from the protein extract. Third, use column
chromatography to fractionate the proteins in the extract and thereby
separate the desired nematode protein from other bacterial proteins.
Because the nematode protein is tagged with multiple histidines, it will
bind to a metal-affinity chromatography matrix. Alternatively, another
matrix can be used, for example an ion-exchange matrix.
D. Yes, the mouse made antibodies against the desired target protein, which
is the band seen only in the gonad lane. No, the serum does not
specifically recognize only one nematode protein; it also recognizes two
proteins found in both gonad and gut cells (Figure A8-5C). To determine
the subcellular location of the target protein you cannot use this serum
because it recognizes additional proteins. To make a cleaner, more specific
antibody preparation, you can create a hybridoma cell line that makes a
unique monoclonal antibody, which is likely to recognize only one
protein. Alternatively, you can make an affinity chromatography matrix
containing the protein of interest, and use affinity chromatography to
purify the desired antibodies out of the mouse serum.
Figure A8-5C
8-6
A. DNA contains only 4 different monomers, and the shape of the double
helix is almost uniform, having only a relatively subtle dependence on the
nucleotide sequence. Protein contains 20 different monomers and the
shapes of proteins differ markedly depending on their amino acid
sequence. Some proteins have long thin rodlike structures and others have
globular structures. All proteins have unique detailed surface contours
with different charge and hydrophobicity properties that allow them to
bind to other molecules and perform their specific unique catalytic or
structural roles.
B. Any specific DNA sequence can be uniquely detected simply by
hybridization to a complementary DNA or RNA. It is easy, inexpensive,
and fast to make a DNA or RNA with any desired sequence (see below).
A protein can be uniquely detected either with a mass spectrometer (MS)
or with a specific antibody. MS equipment costs hundreds of thousands of
dollars and demands highly trained operators. Production of antibodies is
expensive and time-consuming, and usually requires the killing of
laboratory animals.
C. To make many copies of a DNA molecule, we can use DNA cloning or
PCR amplification. PCR requires only two small chemically synthesized
oligonucleotide primers and a small automated reaction that takes a few
hours. DNA cloning requires restriction enzymes, ligase, and a host
plasmid; once a recombinant plasmid has been created, we can make
nearly limitless supplies by growing bacteria harboring the plasmid and
performing a simple standard DNA purification, regardless of nucleotide
sequence. To make many copies of a protein, we can overexpress a gene
from a recombinant expression plasmid, then purify the accumulated
protein away from other proteins with the use of a series of
chromatographic columns. It takes dedicated trial-and-error and patience
to find chromatography conditions that are uniquely suited to purification
of the protein of interest, because each protein behaves differently.
D. Yes, you can unambiguously predict an amino acid sequence from a
cDNA sequence, because each codon specifies a unique amino acid.
However, you cannot unambiguously predict a cDNA sequence from an
amino acid sequence, because most amino acids are specified by more
than one nucleotide triplet, or codon.
8-7
A. You can look in sequence databases compiled from many organisms to try
to find other genes that encode proteins with similar amino acid
sequences. If you find a related protein that has been studied in another
organism, it may suggest a function for Unk1. For example, if Unk1 is
very similar in amino acid sequence to a known protein kinase, then it
probably functions as one.
B. The simplest first experiment is to knock out Unk1 gene expression by
using RNAi and to evaluate the phenotype of the resulting organisms.
Operationally, you can do this by obtaining a bacterial strain harboring a
DNA plasmid that directs the expression of double-stranded RNA with the
same sequence as Unk1. You can feed these bacteria to nematodes, which
will take up the RNA into their cells, where it will direct the degradation
of Unk1 mRNA and alteration of chromatin at the Unk1 gene locus.
C. There are two or more acceptable answers. (1) You can discover the
protein–protein interactions by using a two-hybrid technique in yeast cells.
Unk1 can be fused to a DNA-binding domain (DBD) and used as bait.
Prey molecules can be fished out of a DNA library in which nematode
cDNAs are fused to a gene encoding a yeast transcriptional activation
domain (TA). One yeast strain among thousands that contains both Unk1-
DBD and an Unk1-interacting partner fused to AS can be identified and
isolated because it will transcribe the reporter gene. (2) You can identify
the protein–protein interactions with mass spectrometry (MS) of an Unk1-
containing multiprotein complex. First, Unk1 and its associated proteins
can be partly purified from nematodes. This can be done with an antibody
raised against Unk1 or by fusing a protein “tag” to the Unk1 protein
expressed in nematodes; in both cases, the Unk1 protein can be fished out
of a complex protein mixture with antibody-linked beads that bind Unk1
or its tag. Then the partly purified mixture can be subjected to MS
analysis. Because the genome of nematodes has been sequenced, computer
algorithms can search for matches between (a) the measured mass-to-
charge ratios of peptides from co-purifying proteins and (b) the mass-to-
charge ratios of peptides predicted to arise from particular proteins
encoded by the nematode genome.
D. There are several acceptable answers. (1) In situ hybridization of the Unk1
mRNA can detect the mRNA in different cells at different times of
development. (2) An antibody that specifically binds the Unk1 protein can
be used for immunofluorescence to detect the protein in different cells at
different times of development. (3) A reporter gene can be put under the
control of regulatory sequences from upstream and downstream of the
Unk1 coding sequence, and production of the reporter can be monitored in
animals. For example, a live transgenic animal harboring a GFP reporter
gene controlled by Unk1 regulatory sequences can be monitored with a
microscope to watch where and when the Unk1 gene is turned on.
8-8
A. The haplotype blocks surrounding the lactase gene in lactase-persistent
individuals are much bigger than average haplotype blocks. The
corresponding haplotype blocks in lactose-intolerant individuals are of
average size. (Actually, the haplotype block containing the lactase gene in
lactase-persistent individuals of European descent is 1000–2000 kilobases,
in comparison with an average block of only 2 kilobases! This provides
one of the strongest genomic signatures of evolutionary history in humans
reported so far. When a genetic variant or allele arises that greatly benefits
its carriers and enables them to produce many more viable offspring than
others in the population, this allele will become the most prevalent one in
a population. If it happens relatively quickly, over tens of generations,
then there will few meioses during which crossovers can occur to separate
this allele from the flanking SNPs. Over time, the genetic linkage between
this allele and flanking SNPs will degrade as a result of recombination and
new mutations. Thus, a large and prevalent haplotype block is evidence
for the recent rise of a particular allele under strong positive selective
pressure. It is fascinating that this lactase-persistence trait seems to have
been strongly selected in humans only where and when they domesticated
mammals for dairy farming!)
B. The information provided is insufficient to conclude that the C/T-13910
SNP causes the lactase-persistence trait. A correlation between a SNP and
a trait, by itself, never proves a causal relationship, and this is especially
true if the SNP falls in a large haplotype block.
C. Either Northern blotting or RT-PCR can determine whether an unknown
adult is lactose-intolerant or lactase-persistent, by measuring whether the
lactase mRNA is absent or present, respectively.
D. Lactase persistence is likely to be a dominant trait, because heterozygous
individuals will make some lactase enzyme and will thus be able to digest
milk.
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"Alora, la bien cercada,—tú que estás en par del río,
cercóte el adelantado—una mañana en domingo,
de peones y hombres de armas—el campo bien guarnecido;
con la gran artillería—hecho te había un portillo".
El Canciller, en el Rimado:
Floro:
Adriano:
"Otro escritor que usó mucho del asonante, añade Bello, bien que
no con la constante regularidad del historiador de Matilde, fué
Gofredo de Viterbo en su Panteon, especie de crónica universal,
sembrada de pasajes en verso". Igualmente en la prosa de San
Pedro Damiano, del siglo xi:
Y nótese que sólo hay asonante cada dos versos, quedando sin él
los intermedios, como si fuesen hemistiquios. Á fines del siglo vi ó
principios del vii en el ritmo de San Columbano:
Ápparébit répentína
díes mágna dóminí
fúr obscúra vélut nócte
ímprovísos óccupáns".
La mayor parte de los versos que suenan bien son del mester de
clerezia; por excepción hay alguno que otro del pie de romance, ni
más ni menos que en el Arcipreste de Hita. No puedo yo creer que
si el autor de Mio Cid se hubiera propuesto versificar en romance
tuviera tan toscas orejas que no lo pudiera hacer. Ni de este metro
caótico del Mio Cid pudo salir el romance. Primero, porque los más
de los versos son del metro del mester de clerezia, y de este metro
no salió ni pudo salir el pie de romance, por ser ritmo enteramente
diferente. Segundo, porque de los versos que no suenan y son
verdaderamente caóticos, ni el pie de romance ni metro alguno pudo
nacer. Lo que aquí hay es que M. Pelayo, por seguir la teoría de
Milá, aceptada también por M. Pidal, de que los romances son
fragmentos desprendidos de las gestas, por consecuencia tuvo que
afirmar que el metro del romance salió del metro de las gestas, y ya
que se veía bien claro no serlo, no halló otra solución que la de
atenerse á que "como representan períodos distintos de nuestra
poesía épica, los romances ofrecen ya en estado relativamente fijo y
normal lo que es incierto y caótico en las gestas". Pero ni los
romances se desprendieron de las gestas, ni "el sistema en unos y
otros es sustancialmente el mismo", como no lo es el metro del
mester de clerezia, que es el de Mio Cid, y el metro de los
romances. Son metros sustancialmente diferentes. El pie de
romance es el octosílabo trocaico español, y el metro del mester de
clerezia y de Mio Cid es el septenario yámbico francés; aquél es el
metro del pueblo, éste de los eruditos; aquél el que evolucionó
desde el latín, éste el que los eruditos trajeron de Francia; aquél el
nacional y, por serlo, el despreciado por los eruditos; éste el extraño
y, por serlo, por los eruditos llevado en palmas.
Pero los himnos que los clérigos cantaban todos los días son los
que remedaron en el alejandrino y cuaderna vía. Fueron sus autores
San Gregorio, Prudencio, San Ambrosio y Sedulio, y recogiólos un
tal Hylarius. Casi todos son yámbicos. "Das Metrum der
altfranzösischen Epen—dice Westphall—ist ebenfalls acht und
siebensylbig, hat aber nicht in dem trochäischen, sondern in dem
iambischen Dimetrum (rerum creator omnium) seinen Ursprung,
denn es beginnt nicht mit dem schweren Takttheile, sondern mit der
Anakrusis". Los himnos eclesiásticos eran casi todos yámbicos,
ritmo que cuadra de lleno á la lengua francesa, la cual tiene agudas
las palabras, al revés del castellano, que teniéndolas de ordinario
graves ó llanas, se avenía mejor con el ritmo trocaico. Por lo mismo,
el septenario ó impar concordaba con el francés, y con el castellano
el octonario ó par. Pudieron, pues, los clérigos españoles emplear
en su versificación castellana el metro yámbico, que cantaban
cotidianamente en latín; pero el hecho de que en la época en que
aparecen los primeros monumentos poéticos eruditos en España,
que son los escritos en este metro de catorce sílabas, con los de
siete y de nueve, es cabalmente cuando se deja sentir tan
poderosamente la influencia de los cluniacenses en la corte y en la
iglesia española, hace creer que á ese influjo eclesiástico francés se
deba su empleo. Con los benedictinos de Cluny vinieron á España
las piezas litúrgicas franco-latinas, por ejemplo, el Auto de los Reyes
Magos, tomado del oficio latinizado de alguna ciudad francesa. La
Vida de Santa María Egipciaqua, en versos de nueve sílabas, por lo
ordinario, está tomada de otra obra francesa; el Libro de Apollonio
está en la cuaderna vía ó versos de siete sílabas, y en el mismo
metro se escribió el Mio Cid, que tiene semejanzas con la Chanson
de Roland, que no pueden ser casuales: el obispo francés don
Jerónimo, fogoso como el arzobispo Turpin en el poema francés,
Álvar Fáñez, "diestro brazo" del Cid, como Roland era el "destre
braz" de Carlomagno.
¿Por qué, pues, se niega que hubiese romances antes del siglo xv?
¿Por qué se añade que los romances conocidos del xv son trozos
desprendidos de gestas versificadas en alejandrinos como el Mio
Cid? Lo que de aquí se saca es que el pueblo tenía sus gestas,
largas ó cortas, en romances, que de ellas pasaron trozos á las
Crónicas y que las gestas que conocemos de Mio Cid y Rodrigo y
Alixandre son imitaciones que los clérigos hicieron de las populares,
trayendo del canto eclesiástico y de Francia un nuevo metro erudito,
que, poco á poco, se perfecciona; pero que raras veces deja la liga
del pie de romance que á los clérigos poetas les reteñía por oirlos
en el pueblo, por más que lo menospreciasen. La tan decantada
gesta de los Infantes de Lara, que tenemos prosificada en una de
las Crónicas, son trozos de romances, parecidísimos á los romances
conservados como tales del mismo asunto. No salieron estos
romances de aquellos otros, llamados gesta: son hermanos
gemelos, acaso unos más antiguos que otros, pero nada más.
Pueden verse cotejados romances y gestas en M. Pelayo (Antol.
poet. lír., cast., t. XI, pág. 276). Pero hay más: algunos trozos
parécense á Mio Cid: ¡como que ésta es la única gesta en que
acaso se fundieron varios romances, aunque versificándola el poeta
por el nuevo mester de clerezia. Cuando conserva el pie de
romance resulta un romance verdadero. Y luego dirán que no hubo
romances hasta el siglo xv. Véase este trozo de la llamada gesta de
los de Lara: