Tema 3.

Biología II

Reproducción celular en eucariotas:

Clase 6. Entrecruzamiento y recombinación. Ligamiento y mapa
genético de eucariotas. ADN en organelos.
Replicación del ADN

viernes, mayo 6, 3:02:23 PM

viernes, mayo 6, 3:02:51 PM
Mitosis
Meiosis

viernes, mayo 6, 3:05:45 PM

viernes, mayo 6, 3:06:01 PM
 DNA Recombination
Mechanisms

viernes, mayo 6, 3:06:21 PM

 Recombination

Present in prokaryotic and eukaryotic cells

Only poorly understood

 Why do chromosomes undergo
recombination?
1. include roles in specialized DNA repair systems,
2. specialized activities in DNA replication,
3. regulation of expression of certain genes,
4. facilitation of proper chromosome segregation during
eukaryotic cell division,
5. maintenance of genetic diversity,
6. and implementation of programmed genetic
rearrangements during embryonic development.
 Recombination
ABCDEFGHIJKLMNOPQRSTUVWXYZ
abcdefghijklmnopqrstuvwxyz

ABCDEFGhijklmnoPQRSTUVWXYZ
abcdefgHIJKLMNOpqrstuvwxyz
 Mitotic and meiotic recombination

Recombination can occur both during mitosis and meiosis

Only meiotic recombination serves the important role of
reassorting genes

Mitotic recombination may be important for repair of

mutations in one of a pair of sister chromatids
 Recombination mechanisms
Best studied in yeast, bacteria and phage

Recombination is mediated by the breakage

and joining of DNA strands
 Classes of Recombination
Genetic recombination events fall into at least three general classes.

1. Homologous genetic recombination

(also called general recombination) involves genetic exchanges between any two
DNA molecules (or segments of the same molecule) that share an extended region of
nearly identical sequence. The actual sequence of bases is irrelevant, as long as it is
similar in the two DNAs.

2. In site-specific recombination, the exchanges occur only at a particular DNA

sequence.

3. DNA transposition is distinct from both other classes in that it usually involves a
short segment of DNA with the remarkable capacity to move from one location in a
chromosome to another. These “jumping genes” were first observed in maize in the
1940s by Barbara McClintock. There is in addition a wide range of unusual genetic
rearrangements for which no mechanism or purpose has yet been proposed
 1. Homologous Recombination
Homologous recombination thus
serves at least three identifiable
functions:

(1) it contributes to the repair of

several types of DNA damage;
(2) it provides, in eukaryotic cells, a
transient physical link between
chromatids that promotes the
orderly segregation of chromosomes
at the first meiotic cell division; and
(3) it enhances genetic diversity in a
population.
Figure: The homologous chromosomes of a grasshopper are shown during prophase I
of meiosis. Many points of joining (chiasmata) are evident between the two
homologous pairs of chromatids. These chiasmata are the physical manifestation of
prior homologous recombination (crossing over) events.
• DNA Recombination Is Directed by Specific Enzymes

• Genetic recombination involves:

• endonuclease nicking
• strand displacement
• ligation
• branch migration
• duplex separation to generate the characteristic Holliday
structure (chi form)
 The Holliday model
Two homologous duplexes are aligned

Strand exchange leads to an intermediate with

crossed strands

This branch can move: Branch migration

The branch is resolved by cleavage and sealing

Recombination during Meiosis Is
Initiated with Double-Strand Breaks

Double
strand
break
model

FIGURE 25–31
Double
strand
break
model
 A Chi site or Chi sequence is a short stretch of DNA in
the genome of a bacterium near which homologous
recombination is more likely than expected to occur. Chi
sites serve as stimulators of DNA double-strand break
repair in bacteria, which can arise from radiation or
chemical treatments, or result from replication
fork breakage during DNA replication. The sequence of
the Chi site is unique to each group of closely related
organisms; in E. coli and other enteric bacteria, such as
Salmonella, the sequence is 5'-GCTGGTGG-3'. The
existence of Chi sites was originally discovered in the
genome of bacteriophage lambda, a virus that infects E.
coli, but is now known to occur about 1000 times in the E.
coli genome.

FIGURE 25–31
A Holliday intermediate formed between two bacterial plasmids in vivo, as seen with the electron
microscope. The intermediates are named for Robin Holliday, who first proposed their existence in 1964.
 Para estudio
Recombination during Meiosis Is Initiated with Double-Strand Breaks
A likely pathway for homologous recombination during meiosis has four key features.
1. First, homologous chromosomes are aligned.
2. Second, a double-strand break in a DNA molecule is enlarged by an exonuclease, leaving a
single strand extension with a free 3-hydroxyl group at the broken end (step 1 ).
3. Third, the exposed 3 ends invade the intact duplex DNA, and this is followed by branch
migration (Fig. 25–32) and/or replication to create a pair of crossover structures, called
Holliday junctions (Fig. 25–31a, steps 2 to 4 ).
4. Fourth, cleavage of the two crossovers creates two complete recombinant
products (step 5 ).
In this double-strand break repair model for recombination, the 3 ends are used to initiate the
genetic exchange. Once paired with the complementary strand on the intact homolog, a region of
hybrid DNA is created that contains complementary strands from two different parent DNAs (the
product of step 2 in Fig. 25–31a).
Each of the 3 ends can then act as a primer for DNA replication. The structures thus formed,
Holliday intermediates (Fig. 25–31b), are a feature of homologous genetic recombination
pathways in all organisms.
 Recombination Requires a Host of Enzymes
and Other Proteins

The RecBCD enzyme is both

a helicase that unwinds, or separates
the strands of DNA, and
a nuclease that makes single-
stranded nicks in DNA

The RecBCD enzyme binds to linear DNA at a free (broken) end and moves inward along the double helix, unwinding and
degrading the DNA in a reaction coupled to ATP hydrolysis (Fig. 25–33). The activity of the enzyme is altered when it
interacts with a sequence referred to as chi, (5)GCTGGTGG. From that point, degradation of the strand with a 3 terminus
is greatly reduced, but degradation of the 5-terminal strand is increased. This process creates a single-stranded DNA with
a 3 end, which is used during subsequent steps in recombination (Fig. 25–31).
Action of E. coli proteins in branch
migration and resolution of Holliday
structures
viernes, mayo 6, 3:30:17 PM
 Recombination between homologous
DNA sites
Recombination provides a means by which a genome can
change to generate new combinations of genes

Homologous recombination allows for the exchange of blocks

of genes between homologous chromosomes and thereby is
a mechanism for generating genetic diversity

Recombination occurs randomly between two homologous

sequences and the frequency of recombination between two
sites is proportional to the distance between the sites
 Classes of Recombination
Genetic recombination events fall into at least three general classes.

1. Homologous genetic recombination

(also called general recombination) involves genetic exchanges between any two
DNA molecules (or segments of the same molecule) that share an extended region of
nearly identical sequence. The actual sequence of bases is irrelevant, as long as it is
similar in the two DNAs.

2. In site-specific recombination, the exchanges occur only at a particular DNA

sequence.

3. DNA transposition is distinct from both other classes in that it usually involves a
short segment of DNA with the remarkable capacity to move from one location in a
chromosome to another. These “jumping genes” were first observed in maize in the
1940s by Barbara McClintock. There is in addition a wide range of unusual genetic
rearrangements for which no mechanism or purpose has yet been proposed
 2.Site specific recombination
Viruses and transposable elements often integrate their
genomes into the host chromosome

Site specific recombination is used by both eukaryotes

and prokaryotes to regulate gene expression and to
increase the organisms genetic range
 Site specific recombination
FIGURE 25–38 A site-specific recombination reaction. (a)
The reaction is carried out within a tetramer of identical
subunits. Recombinase subunits bind to a specific sequence,
often called simply the recombination site.
1 One strand in each DNA is cleaved at particular points
within the sequence. The nucleophile is the OH group of an
active-site Tyr residue, and the product is a covalent
phosphotyrosine link between protein and DNA.
2 The cleaved strands join to new partners, producing
a Holliday intermediate.
Steps 3 and 4 complete the reaction by a process similar to
the first two steps.
The original sequence of the recombination site is
regenerated after recombining the DNA flanking the site.
These steps occur within a complex of multiple recombinase
subunits that sometimes includes other proteins.
 Site specific recombination

Each site-specific recombination system consists of an enzyme called a recombinase and a

short (20 to 200 bp), unique DNA sequence where the recombinase acts (the recombination
site). A separate recombinase recognizes and binds to each of two recombination sites on two
different DNA molecules or within the same DNA. One DNA strand in each site is cleaved at a
specific point within the site, and the recombinase becomes covalently linked to the DNA at
the cleavage site through a phosphotyrosine (or phosphoserine) bond (step 1 ). The transient
protein-DNA linkage preserves the phosphodiester bond that is lost in cleaving the DNA, so
high-energy cofactors such as ATP are unnecessary in subsequent steps. The cleaved DNA
strands are rejoined to new partners to form a Holliday intermediate, with new
phosphodiester bonds created at the expense of the protein-DNA linkage (step 2 ). To
complete the reaction, the process must be repeated at a second point within each of the two
recombination sites (steps 3 and 4 ). In some systems, both strands of each recombination
site are cut concurrently and rejoined to new partners without the Holliday intermediate. The
exchange is always reciprocal and precise, regenerating the recombination sites when the
reaction is complete. We can view a recombinase as a site-specific endonuclease and ligase in
one package.
 Site specific recombination

FIGURE 25–39 Effects of site-specific recombination. The outcome of site-specific recombination depends on the
location and orientation of the recombination sites (red and green) in a double-stranded DNA molecule. Orientation here
(shown by arrowheads) refers to the order of nucleotides in the recombination site, not the 5n3 direction.
(a) Recombination sites with opposite orientation in the same DNA molecule. The result is an inversion. (b)
Recombination sites with the same orientation, either on one DNA molecule, producing a deletion, or on two DNA
molecules, producing an insertion.
 Ejemplo de recombinación

The immune system is a system of many

biological structures and processes within Immune system
an organism that protects against disease. To
function properly, an immune system must
detect a wide variety of agents, known
as pathogens, from viruses to parasitic worms,
and distinguish them from the organism's own
healthy tissue. In many species, the immune
system can be classified into subsystems, such
as the innate immune system versus
the adaptive immune system, or humoral
immunity versus cell-mediated immunity.
Humoral immunity, also called the antibody-mediated
beta cellularis immune system, is the aspect
of immunity that is mediated by macromolecules (as
opposed to cell-mediated immunity) found in extracellular
fluids such as secreted antibodies, complement
proteins and certain antimicrobial peptides. Humoral
immunity is so named because it involves substances
found in the humours, or body fluids.
immunoglobulin
immunoglobulin
 Site specific recombination

When phage DNA enters an E. coli

cell, a complex series of regulatory
events commits the DNA to one of
two fates. The DNA either
replicates and produces more
bacteriophages (destroying the
host cell) or integrates into the
host chromosome, replicating
passively along with the
chromosome for many cell
generations. Integration is
accomplished by a phage-encoded
recombinase ( integrase) that acts
at recombination sites on the
phage and bacterial DNAs—at
attachment sites attP and attB,
respectively (Fig. 25–40).
FIGURE 25–40 Integration and excision of bacteriophage DNA at the chromosomal target site. The attachment site
on the phage DNA (attP) shares only 15 bp of complete homology with the bacterial site (attB) in the region of the
crossover. The reaction generates two new attachment sites (attR and attL) flanking the integrated phage DNA.
The recombinase is the integrase (or INT protein). Integration and excision use different attachment sites and different
auxiliary proteins. Excision uses the proteins XIS, encoded by the bacteriophage, and FIS, encoded by the bacterium. Both
reactions require the protein IHF (integration host factor), encoded by the bacterium.
 Classes of Recombination
Genetic recombination events fall into at least three general classes.

1. Homologous genetic recombination

(also called general recombination) involves genetic exchanges between any two
DNA molecules (or segments of the same molecule) that share an extended region of
nearly identical sequence. The actual sequence of bases is irrelevant, as long as it is
similar in the two DNAs.

2. In site-specific recombination, the exchanges occur only at a particular DNA

sequence.

3. DNA transposition is distinct from both other classes in that it usually involves a
short segment of DNA with the remarkable capacity to move from one location in a
chromosome to another. These “jumping genes” were first observed in maize in the
1940s by Barbara McClintock. There is in addition a wide range of unusual genetic
rearrangements for which no mechanism or purpose has yet been proposed
 DNA Transposition

recombination that allows the movement of transposable elements, or transposons. These segments of DNA,
found in virtually all cells, move, or “jump,” from one place on a chromosome (the donor site) to another on the
same or a different chromosome (the target site). DNA sequence homology is not usually required for this
movement, called transposition; the new location is determined more or less randomly. Insertion of a
transposon in an essential gene could kill the cell, so transposition is tightly regulated and usually very
infrequent. Transposons are perhaps the simplest of molecular parasites, adapted to replicate passively within
the chromosomes of host cells. In some cases they carry genes that are useful to the host cell, and thus exist in
a kind of symbiosis with the host
 Classes of Transposons
Bacteria have two classes of transposons.
1. Simple transposons
Insertion sequences contain only the sequences required for transposition and the
genes for proteins (transposases) that promote the process.
2. Complex transposons contain one or more genes in addition to those needed
for transposition. These extra genes might, for example, confer resistance to
antibiotics and thus enhance the survival chances of the host cell. The spread of
antibiotic-resistance elements among disease-causing bacterial populations that is
rendering some antibiotics ineffectual is mediated in part by transposition.
Bacterial transposons vary in structure, but most have short repeated sequences at
each end that serve as binding sites for the transposase. When transposition
occurs, a short sequence at the target site (5 to 10 bp) is duplicated to form an
additional short repeated sequence that flanks each end of the inserted transposon
(Fig. 25–42). These duplicated segments result from the cutting mechanism used
to insert a transposon into the DNA at a new location.
FIGURE 25–42 Duplication
of the DNA sequence at a
target site
when a transposon is
inserted. The duplicated
sequences are shown
in red. These sequences are
generally only a few base
pairs long, so their size
(compared with that of a
typical transposon) is
greatly exaggerated in this
drawing.
 GENETIC LINKAGE= Ligamiento genético
• Genes on nonhomologous chromosomes assort independently during meiosis.
• Genes on the same chromosome are said to exhibit linkage and are called linked
genes.
• Linked genes, and hence the phenotypic characters they control, are inherited
together because they are located on the same chromosome.
• Modern understanding of genetic linkage came from the work of Thomas
Morgan. Morgan showed that two recessive genes in Drosophila melanogaster;
white eye (w) and miniature wing (m) are X-linked.
Concepto de Ligamiento: por definición, se dice que dos
loci están ligados cuando se encuentran situados sobre el
mismo cromosoma. Todos aquellos loci que se
encuentran situados sobre el mismo cromosoma forman
un Grupo de Ligamiento.
Cuanto más alejados están entre sí dos loci ligados ( A,a y C,c) más probable es que se dé
sobrecruzamiento entre ellos, cuanto más cerca están entre sí dos loci ligados (A,a y B,b) menos probable
es que se dé sobrecruzamiento entre ambos.

Dos loci ligados pueden estar en Fase de Acoplamiento AB/ab (los dos alelos dominantes sobre el mismo
cromosoma, y los dos recesivos sobre el cromosoma homologo) o en Fase de Repulsión Ab/aB (un alelo
dominante y otro recesivo sobre cada cromosoma).
• Morgan crossed a female with white eyes and miniature wings
(wm/wm) with a wild-type male (red eyes and large wings) (w+m+/Y).
• The F1 males were white-eyed and had miniature wings (genotype
wm/Y) while the females were wild-type for both the eye colour and
wing size (genotype w+m+/wm).
• The F1 flies were interbred (F1 x F1) and the resulting F2 flies were
analyzed.
• In the F2, the flies fell into four classes: (a) white eyes, miniature wings
were 359 females and 391 males, (b) wild-type (red eyes, large wings)
were 439 females and 352 males, (c) white eyes, large wings were 218
females and 237 males, (d) red eyes, miniature wings were 235 females
and 210 males.
• In the F2 the most frequent phenotypic classes in both sexes were the
grandparental phenotypes (class a and b).
• The genotypes or phenotypes of the original parents are referred to as
parental classes.
• There were also flies with the nonparental phenotypic combinations
(class c and d). Nonparental combinations of linked genes are called
recombinants.
• To explain for the recombinants he proposed that in meiosis, exchange
of genes had occurred between the two X chromosomes of the F1
females. Since males are hemizygous, no such genetic exchange
occurred between X and Y chromosomes.
• The production of recombinants results from physical exchanges
between homologous chromosomes during meiotic prophase I.
 • Crossing-over is the reciprocal exchange of
chromosome parts at corresponding
Crossing-over positions along homologous chromosomes
by symmetrical breakage and rejoining.
Crossing-over is the event that leads to
genetic recombination between linked
genes.
• Morgan’s conclusion was that during
meiosis, alleles of some genes assort
together because they lie near each other
on the same chromosome.
• In many cases, two alleles inherited from
one parent show a strong tendency to stay
together as do those from the other
parent. This phenomenon is called linkage.
• The closer two genes are on the
chromosome, the more likely they are to
remain together during meiosis.
Construction of genetic maps
• Genetic mapping is the process that uses
genetic experiment to determine the
relative position of genes on
chromosomes in eukaryotic organisms.
• A genetic map gives the order in which
genes are arranged along a chromosome
and the spacing between them on the
same chromosome.
• The map distance between two genes is
based on the frequency of
recombination between the two genes.
The recombination frequency is an
approximation of the frequency of
crossovers between the two genes.
• The frequency of crossing-over, and
hence recombinants, for linked genes is
characteristic of the gene pairs involved
e.g. for the white eye (w) and miniature
wing (m) which are X-linked the
frequency of crossing-over is 36.9%
 Organelle DNAs
Although the vast majority of DNA in most eukaryotes is found in the nucleus, some DNA is
present within the mitochondria of animals, plants, and fungi and within the chloroplasts of
plants. These organelles are the main cellular sites for ATP formation, during oxidative
phosphorylation in mitochondria and photosynthesis in chloroplasts (Chapter 16). Many lines
of evidence indicate that mitochondria and chloroplasts evolved from bacteria that were
endocytosed into ancestral cells containing a eukaryotic nucleus,
forming endosymbionts. Over evolutionary time, most of the bacterial genes encoding
components of the present-day organelles were transferred to the nucleus. However,
mitochondria and chloroplasts in today’s eukaryotes retain circular DNAs encoding proteins
essential for organellar function as well as the ribosomal and transfer RNAs required for
their translation. Thus eukaryotic cells have multiple genetic systems: a predominant nuclear
system and secondary systems with their own DNA in the mitochondria and chloroplasts.

http://www.ncbi.nlm.nih.gov/books/NBK21574/
Molecular Cell Biology. 4th edition.
viernes, mayo 6, 7:15:04 PM
 Mitochondria Contain Multiple mtDNA Molecules

Individual mitochondria are large

enough to be seen under the light
microscope and even the
mitochondrial DNA (mtDNA) can
be detected by fluorescence
microscopy. The mtDNA is located
in the interior of the mitochondrion,
the region known as the matrix.
Genes in mtDNA Exhibit Cytoplasmic Inheritance and Encode rRNAs, and
Some Mitochondrial Proteins

Studies of mutants in yeasts and other single-celled organisms

first indicated that mitochondria exhibit cytoplasmic
inheritance and thus must contain their own genetic system
(Figure 9-43). For instance, petite yeast mutants exhibit
structurally abnormal mitochondria and are incapable of
oxidative phosphorylation. As a result, petite cells grow more
slowly than wild-type yeasts and form smaller colonies (hence
the name “petite”). Genetic crosses between different (haploid)
yeast strains showed that the petite mutation does not segregate
with any known nuclear gene or chromosome. In later studies,
most petite mutants were found to contain deletions of mtDNA.
Petite-strain mitochondria are defective in oxidative phosphorylation due to a deletion in
mtDNA. (a) Haploid cells fuse to produce a diploid cell that undergoes meiosis, during which
random segregation of parental chromosomes and mitochondria containing mtDNA occurs. Since
yeast normally contain ≈50 mtDNA molecules per cell, all products of meiosis usually contain
both normal and petite mtDNAs and are capable of respiration. (b) As these cells grow and divide
mitotically, the cytoplasm (including the mitochondria) is randomly distributed to the daughter
cells. Occasionally, a cell is generated that contains only defective petite mtDNA and yields a
petite colony. Thus formation of such petite cells is independent of any nuclear genetic marker.
 The Size and Coding Capacity of mtDNA Vary
Considerably in Different Organisms

Surprisingly, the size of the mtDNA, the number and nature of

the proteins it encodes, and even the mitochondrial genetic
code itself, vary greatly among different organisms. Human
mtDNA, a circular molecule that has been completely sequenced,
is among the smallest known mtDNAs, containing
16,569 base pairs (Figure 9-44). It encodes the two rRNAs found
in mitochondrial ribosomes and the 22 tRNAs used to translate
mitochondrial mRNAs. Human mtDNA has 13 sequences that
begin with an ATG (methionine) codon, end with a stop codon,
and are long enough to encode a polypeptide of more than 50
amino acids; all of the possible proteins encoded by these open
reading frames have been identified. Mammalian mtDNA, in
contrast to nuclear DNA, lacks introns and contains no long
noncoding sequences.
 Mitochondrial Genetic Codes Differ from the Standard Nuclear Code

The genetic code used in animal and fungal mitochondria is different from the standard code used in all prokaryotic and
eukaryotic nuclear genes; remarkably, the code even differs in mitochondria from different species (Table 9-4). Why and how
this phenomenon happened during evolution is mysterious. UGA, for example, is normally a stop codon, but is read as
tryptophan by human and fungal mitochondrial translation systems; however, in plant mitochondria, UGA is still a stop
codon. AGA and AGG, the standard nuclear codons for arginine also code for arginine in fungal and plant mtDNA, but they
are stop codons in mammalian mtDNA and serine codons inDrosophila mtDNA.
Mitochondria
Codon Standard Code: Mammals Drosophila Neurospora Yeasts Plants
Nuclear-Encoded
Proteins
UGA Stop Trp Trp Trp Trp Stop
AGA, AGG Arg Stop Ser Arg Arg Arg
AUA Ile Met Met Ile Met Ile
AUU Ile Met Met Met Met Ile
CUU, CUC, CUA, Leu Leu Leu Leu Thr Leu
CUG

Uso de codones
 Mutations in Mitochondrial DNA Cause Several Genetic Diseases in Man

The severity of disease caused by a mutation in mtDNA

depends on the nature of the mutation and on the
proportion of mutant and wild-type DNAs present in a
particular cell type. Generally, when mutations in mtDNA
are found, cells contain mixtures of wild-type and mutant
mtDNAs — a condition known as heteroplasmy. Each
time a mammalian somatic or germ-line cell divides, the
mutant and wild-type mtDNAs will segregate randomly
into the daughter cells, as occurs in yeast cells (see Figure
9-43). Thus, the mtDNA genotype fluctuates from one
generation and from one cell division to the next, and can
drift toward predominantly wild-type or predominantly
mutant mtDNAs. Since all enzymes for the replication
and growth of mitochondria, such
as DNA and RNA polymerases, are imported from
the cytosol, a mutant mtDNA should not be at a
“replication disadvantage”; mutants that involve large
deletions of mtDNA might even be at a selective
advantage in replication.
Órganos Afectación
Cerebro Retraso en el desarrollo, retardo mental, demencia, convulsiones, desórdenes neuro-
psiquiátricos, parálisis cerebral atípica, migrañas, infartos.
Nervios Debilidad, dolor nueropático, ausencia de reflejos, problemas gastrointestinales (reflujo
gastroesofágeo, vaciado gástrico retrasado, constipación, pseudo obstrucción), desmayos,
ausencia o exceso de sudor relacionados con problemas de regulación de la temperatura.

Músculos Debilidad, hipotonía, calambres, dolor muscular.

Riñones Desgasta proximal renal tubular que provoca pérdida de proteínas, magnesio, fósforo, calcio
y otros electrolitos.
Corazón Defectos en los conductos cardiacos (bloqueos del corazón), cardiomiopatía.
Hígado Hipoglicemia (niveles de azúcar bajos en la sangre), falla del hígado.
Ojos Pérdida de visión y ceguera.
Oídos Pérdida auditiva y sordera.
Páncreas Diabetes y falla pancreatítica exocrina (incapacidad para generar encimas digestivas).
Sistémico Incapacidad para subir de peso, corta estatura, fatiga, problemas respiratorios incluyendo
sofocamientos intermitentes.

martes, noviembre 24, 9:32:02 AM

 Chloroplasts Contain Large Circular DNAs Encoding More Than a Hundred Proteins

the structure of chloroplasts is similar in many respects to that of mitochondria. Like mitochondria, chloroplasts
contain multiple copies of the organellar DNA and ribosomes, which synthesize some chloroplast-encoded
proteins using the “standard” genetic code. Other chloroplast proteins are fabricated on cytosolic ribosomes and
are incorporated into theorganelle after translation.

martes, noviembre 24, 9:32:32 AM

Chloroplast DNAs are circular molecules of 120,000 – 160,000
bp, depending on the species. The complete sequences of
several chloroplast DNAs have been determined, including
those from liverwort (121,024 bp) and tobacco (155,844 bp).
The liverwort chloroplast genome has two inverted repeats, each
consisting of 10,058 bp, that contain the rRNA genes and a few
other duplicated genes. Despite the difference in size, the
overall organization and gene composition of the liverwort and
tobacco DNAs are very similar; the size differential is due
primarily to the length of the inverted repeat in which some
genes are duplicated.
Of the ≈120 genes in chloroplast DNA, about 60 are involved
in RNA transcription and translation, including genes for
rRNAs, tRNAs, RNA polymerase subunits, and ribosomal
proteins. About 20 genes encode subunits of the chloroplast
photosynthetic electron transport complexes and the
F0F1ATPase complex. Also encoded in the
chloroplast genome is the larger of the two subunits of ribulose
1,5-bisphosphate carboxylase, which is involved in the fixation
of carbon dioxide duringphotosynthesis.
Reflecting the endosymbiotic origin of chloroplasts, some
regions of chloroplast DNA are strikingly similar to those of the
DNA of present-day bacteria. For instance, chloroplast DNA
encodes four subunits of RNA polymerase that are highly
homologous to the subunits of E. coliRNA polymerase. One
segment of chloroplast DNA encodes eight proteins that are
homologous to eight E. coli ribosomal proteins; the order of
these genes is the same in the two DNAs.
Liverwort (briófitas) chloroplast DNA has some genes that are
not detected in the larger tobacco chloroplast DNA, and vice
versa. Since the two types of chloroplasts contain virtually the
same set of proteins, these data suggest that some genes are
present in the chloroplast DNA of one species and in the nuclear
DNA of the other, indicating that some exchange of genes
between chloroplast and nucleushas occurred during evolution.
 Clase 6.
Entrecruzamiento y recombinación.
Ligamiento y mapa genético de eucariotas.
ADN en organelos.