Transcripción Alberts 2014 PDF

Alberts Johnson Lewis Morgan Raff Roberts Walter
Molecular Biology
of the Cell
Sixth Edition
Chapter 6
How Cells Read the Genome:
From DNA to Protein
Copyright Garland Science 2015
CHAPTER CONTENTS
FROM DNA TO RNA
FROM RNA TO PROTEIN
THE RNA WORLD AND THE ORIGINS OF LIFE
Introduction
Introduction
FROM DNA TO RNA

Introduction
Introduction
FROM DNA TO RNA

RNA Molecules Are Single-Stranded
FROM DNA TO RNA

Transcription Produces RNA Complementary to
One Strand of DNA
Transcription Produces RNA Complementary to One Strand of DNA
FROM DNA TO RNA

RNA Polymerases Carry Out Transcription
FROM DNA TO RNA

Cells Produce Different Categories of RNA
Molecules
Cells Produce Different Categories of RNA Molecules
FROM DNA TO RNA

Signals Encoded in DNA Tell RNA Polymerase
Where to Start and Stop
Signals Encoded in DNA Tell RNA Polymerase Where to Start and

Stop

Stop

Stop

Stop
FROM DNA TO RNA

Transcription Start and Stop Signals Are
Heterogeneous in Nucleotide Sequence
Transcription Start and Stop Signals Are Heterogeneous in

Nucleotide Sequence

Nucleotide Sequence

Nucleotide Sequence

Nucleotide Sequence

Nucleotide Sequence
FROM DNA TO RNA

Transcription Initiation in Eukaryotes Requires
Many Proteins
Transcription Initiation in Eukaryotes Requires Many Proteins
Las 3 RNA polimerasas son estructuralemente similares entre ellas (y con la RNA polimerasa bacteriana)
comparten subunidades pero transcriben diferente tipo de genes.
La RNA polimerasa II tiene muchas similitudes estructurales con la RNA polimerasa bacteriana, pero tambin
existen diferencias importantes:
1- La RNA polimerasa bacteriana requiere solo de una protena adicional para la iniciacin de la transcripcin,
la RNA polimerasa II requiere de muchas protenas adicionales,
llamadas factores de transcripcin generales.
2- La iniciacin de la transcripcin en eucariotes trata con el empacamiento de DNA en nucleosomas y
en cromatina, una estructura altamente organizada, ausente en el cromosoma bacteriano.
Transcription Initiation in Eukaryotes Requires Many Proteins
FROM DNA TO RNA

RNA Polymerase II Requires a Set of General
Transcription Factors
RNA Polymerase II Requires a Set of General Transcription Factors
Los factores de transcripcin generales

ayudan a posicionar a la RNA
polimerasa correctamente sobre el
promotor, ayudan a separar la doble
hlice para que comience la
transcripcin y a liberar a la RNA
polimerasa del promotor al modo de
elongacin, una vez que inici la
transcripcin. Se dice que los factores
son generales porque se requieren en
casi todos los promotores de la RNApol
II.
Secuencias consenso que se han encontrado cerca del inicio de la transcripcin
FROM DNA TO RNA

Polymerase II Also Requires Activator, Mediator,
and Chromatin-Modifying Proteins
La transcripcin por la RNA pol II en una clula eucarionte es ms compleja y

requiere muchas ms protenas.
1- requiere de protenas regulatorias - activadores transcripcionales, los cuales
deben unirse a secuencias de DNA especficas y ayudan a atraer la RNA pol II
al punto de inicio de la transcripcin
2- la iniciacin de la transcripcin en eucariontes in vivo requiere de un
complejo protico - mediador, que permite que las protenas activadoras se
comuniquen con la RNA pol II y con los factores de transcripcin generales.
3- la iniciacin de la transcripcin en eucariontes requiere el reclutamiento local
de las enzimas modificadoras de la cromatina, incluyendo los complejos
remodeladores de la cromatina y las enzimas modificadoras de las histonas.
Polymerase II Also Requires Activator, Mediator, and ChromatinModifying Proteins
FROM DNA TO RNA

Transcription Elongation in Eukaryotes Requires
Accessory Proteins
FROM DNA TO RNA

Transcription Creates Superhelical Tension
FROM DNA TO RNA

Transcription Elongation in Eukaryotes Is Tightly
Coupled to RNA Processing
Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA

Processing

Processing

Processing

Processing

Processing

Processing
la transcripcin en eucariontes est acoplada al

procesamiento del RNA
El factor de transcripcin TFIIH fosforila el

CTD de la RNA polimerasa II, indicando la
transicin de la iniciacin a la elongacin. Las
protenas necesarias para el capping, para el
procesamiento y para la poliadenilacin del pre
RNAm reconocen las diferentes formas
fosforiladas del CTD de la RNA polimerasa II.
De lo que se trata es que las protenas
encargadas del capping, de procesamiento del
pre RNAm y de la poliadenilacin del RNAm
se acerquen a sus sitios de accin en el
RNAm precursor conforme se transcriben
durante la elongacin.

Processing

Processing
Conforme la polimerasa transcribe, va cargando protenas de procesamiento del preRNAm en su cola (CTD)
las cuales va a transferir al RNAm naciente. La protena de capping se une primero al CTD de la RNA polimerasa
cuando est fosforilada en la ser5 del hepta-repetido, una vez que la iniciacin de la transcripcin est avanzada.
Esto asegura que la molcula de RNA se modifique eficientemente en el 5.
Posteriormente, el CTD de la polimerasa se fosforila en la ser2 por una kinasa asociada con las protenas de elongacin,
y eventualmente se desfosforila en la ser5. Estas modificaciones en el CTD de la polimerasa van a atraer otras protenas
de procesamiento para que acten en el RNAm conforme va emergiendo de la RNA polimerasa.
Sin embargo son muchas las enzimas que procesan el RNAm, y no todas viajan con la RNA polimerasa.
Pero una vez que alguna protena crtica del procesamiento se ha transferido a la molcula de RNAm,
va a atraer al resto del complejo proteco para que procese la molcula de RNA.
FROM DNA TO RNA

RNA Capping Is the First Modification of Eukaryotic
Pre-mRNAs
RNA Capping Is the First Modification of Eukaryotic Pre-mRNAs
3 enzimas, que actan sucesivamente, realizan la reaccin de capping.

Una fosfatasa elimina un fosfato del extremo 5 nasciente del RNA,
una guanidiltransferasa aade un GMP en 5-5,
y una metil transferasa metila la guanosina.
Las 3 se unen al CTD fosforilado en la ser5,
y por lo tanto estn listas para modificar el extremo 5 naciente
al momento que emerge de la polimerasa.
El cap 5 metilado significa un extremo 5 de un RNAm,

y esto le permite a la clula distinguir entre otros tipos de molculas
de RNA presentes en la clula. La RNA polI y la RNA pol III no
producen transcritos con cap, en parte porque no tienen CTD.
Cual es la funcin del cap en el extremo 5 del RNAm?

En el ncleo, el cap del 5 del RNAm se une a un complejo protico (CBC)
que le ayuda al mensajero a procesarse y exportarse adecuadamente.
Tambin el cap tiene una funcin en la traduccin.
FROM DNA TO RNA

RNA Splicing Removes Intron Sequences from
Newly Transcribed Pre-mRNAs
RNA Splicing Removes Intron Sequences from Newly Transcribed

Pre-mRNAs

Pre-mRNAs

Pre-mRNAs

Pre-mRNAs

Pre-mRNAs

Pre-mRNAs

Pre-mRNAs
FROM DNA TO RNA

Nucleotide Sequences Signal Where Splicing
Occurs
Nucleotide Sequences Signal Where Splicing Occurs
FROM DNA TO RNA

RNA Splicing Is Performed by the Spliceosome
El splicing de RNA est hecho por el

spliceosoma
Los pasos crticos en el procesamiento de RNA estn hechos por
molculas de RNA- no por protenas. Las molculas de RNA reconocen las

secuencias de nucletidos que especifican los sitios de procesamiento y
tambin participan en la qumica del procesamiento (la catlisis).
Son molculas de RNA pequeas (menos de 200nt) y 5 (U1, U2, U4, U5 y U6)
son las que estn involucradas en el tipo principal de splicing del pre-RNAm.
Son los snRNAs (RNAs pequeos nucleares), y cada uno forma un complejo con
al menos 7 subunidades proticas, y forman un snRNP (ribonucleoprotena
pequea nuclear). Estas snRNPs forman la mdula de spliceosoma, el complejo
de RNA y protenas que realiza el splicing.
FROM DNA TO RNA

The Spliceosome Uses ATP Hydrolysis to Produce a
Complex Series of RNARNA Rearrangements
The Spliceosome Uses ATP Hydrolysis to Produce a Complex Series of

RNARNA Rearrangements
FROM DNA TO RNA

Other Properties of Pre-mRNA and Its Synthesis
Help to Explain the Choice of Proper Splice Sites
Other Properties of Pre-mRNA and Its Synthesis Help to Explain the

Choice of Proper Splice Sites






FROM DNA TO RNA

Chromatin Structure Affects RNA Splicing
FROM DNA TO RNA

RNA Splicing Shows Remarkable Plasticity
RNA Splicing Shows Remarkable Plasticity
FROM DNA TO RNA

Spliceosome-Catalyzed RNA Splicing Probably
Evolved from Self-splicing Mechanisms
FROM DNA TO RNA

RNA-Processing Enzymes Generate the 3 End of
Eukaryotic mRNAs
RNA-Processing Enzymes Generate the 3 End of Eukaryotic mRNAs
FROM DNA TO RNA

Mature Eukaryotic mRNAs Are Selectively
Exported from the Nucleus
Mature Eukaryotic mRNAs Are Selectively Exported from the Nucleus
FROM DNA TO RNA

Noncoding RNAs Are Also Synthesized and
Processed in the Nucleus
Noncoding RNAs Are Also Synthesized and Processed in the Nucleus
FROM DNA TO RNA

The Nucleolus Is a Ribosome-Producing Factory
FROM DNA TO RNA

The Nucleus Contains a Variety of Subnuclear
Aggregates
The Nucleus Contains a Variety of Subnuclear Aggregates
FROM RNA TO PROTEIN

An mRNA Sequence Is Decoded in Sets of Three
Nucleotides
An mRNA Sequence Is Decoded in Sets of Three Nucleotides
An mRNA Sequence Is Decoded in Sets of Three Nucleotides
FROM RNA TO PROTEIN

tRNA Molecules Match Amino Acids to Codons in
mRNA
tRNA Molecules Match Amino Acids to Codons in mRNA
FROM RNA TO PROTEIN

tRNAs Are Covalently Modified Before They Exit
from the Nucleus
tRNAs Are Covalently Modified Before They Exit from the Nucleus
tRNAs Are Covalently Modified Before They Exit from the Nucleus
FROM RNA TO PROTEIN

Specific Enzymes Couple Each Amino Acid to Its
Appropriate tRNA Molecule
Specific Enzymes Couple Each Amino Acid to Its Appropriate tRNA

Molecule

Molecule

Molecule

Molecule

Molecule
FROM RNA TO PROTEIN

Editing by tRNA Synthetases Ensures Accuracy
FROM RNA TO PROTEIN

Amino Acids Are Added to the C-terminal End of a
Growing Polypeptide Chain
Amino Acids Are Added to the C-terminal End of a Growing

Polypeptide Chain
FROM RNA TO PROTEIN

The RNA Message Is Decoded in Ribosomes
FROM RNA TO PROTEIN

Elongation Factors Drive Translation Forward and
Improve Its Accuracy
Elongation Factors Drive Translation Forward and Improve Its

Accuracy

Accuracy

Accuracy

Accuracy
FROM RNA TO PROTEIN

Many Biological Processes Overcome the Inherent
Limitations of Complementary Base-Pairing
FROM RNA TO PROTEIN

Accuracy in Translation Requires an Expenditure of
Free Energy
FROM RNA TO PROTEIN

The Ribosome Is a Ribozyme
FROM RNA TO PROTEIN

Nucleotide Sequences in mRNA Signal Where to
Start Protein Synthesis
Nucleotide Sequences in mRNA Signal Where to Start Protein

Synthesis

Synthesis

Synthesis

Synthesis
FROM RNA TO PROTEIN

Stop Codons Mark the End of Translation
Stop Codons Mark the End of Translation
FROM RNA TO PROTEIN

Proteins Are Made on Polyribosomes
FROM RNA TO PROTEIN

There Are Minor Variations in the Standard
Genetic Code
There Are Minor Variations in the Standard Genetic Code
FROM RNA TO PROTEIN

Inhibitors of Prokaryotic Protein Synthesis Are
Useful as Antibiotics
Inhibitors of Prokaryotic Protein Synthesis Are Useful as Antibiotics
Inhibitors of Prokaryotic Protein Synthesis Are Useful as Antibiotics
FROM RNA TO PROTEIN

Quality Control Mechanisms Act to Prevent
Translation of Damaged mRNAs
Quality Control Mechanisms Act to Prevent Translation of Damaged

mRNAs
FROM RNA TO PROTEIN

Some Proteins Begin to Fold While Still Being
Synthesized
Some Proteins Begin to Fold While Still Being Synthesized
FROM RNA TO PROTEIN

Molecular Chaperones Help Guide the Folding of
Most Proteins
FROM RNA TO PROTEIN

Cells Utilize Several Types of Chaperones
FROM RNA TO PROTEIN

Exposed Hydrophobic Regions Provide Critical
Signals for Protein Quality Control
Exposed Hydrophobic Regions Provide Critical Signals for Protein

Quality Control
FROM RNA TO PROTEIN

The Proteasome Is a Compartmentalized Protease
with Sequestered Active Sites
The Proteasome Is a Compartmentalized Protease with Sequestered

Active Sites

Active Sites

Active Sites

Active Sites

Active Sites

Active Sites

Active Sites
FROM RNA TO PROTEIN

Many Proteins Are Controlled by Regulated
Destruction
Many Proteins Are Controlled by Regulated Destruction
FROM RNA TO PROTEIN

There Are Many Steps From DNA to Protein
There Are Many Steps From DNA to Protein
THE RNA WORLD AND THE

ORIGINS OF LIFE
Introduction
Introduction

ORIGINS OF LIFE
Single-Stranded RNA Molecules Can Fold into
Highly Elaborate Structures
Single-Stranded RNA Molecules Can Fold into Highly Elaborate

Structures

Structures

Structures

Structures

Structures

Structures

ORIGINS OF LIFE
RNA Can Both Store Information and Catalyze
Chemical Reactions
RNA Can Both Store Information and Catalyze Chemical Reactions

ORIGINS OF LIFE
How Did Protein Synthesis Evolve?

ORIGINS OF LIFE
All Present-Day Cells Use DNA as Their Hereditary
Material
All Present-Day Cells Use DNA as Their Hereditary Material
PROBLEMS
PROBLEMS
PROBLEMS
PROBLEMS
PROBLEMS
PROBLEMS

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Alberts Johnson Lewis Morgan Raff Roberts Walter

Copyright Garland Science 2015

FROM DNA TO RNA

FROM DNA TO RNA

RNA Molecules Are Single-Stranded

RNA Molecules Are Single-Stranded

RNA Molecules Are Single-Stranded

RNA Molecules Are Single-Stranded

RNA Molecules Are Single-Stranded

RNA Molecules Are Single-Stranded

RNA Molecules Are Single-Stranded

RNA Molecules Are Single-Stranded

RNA Molecules Are Single-Stranded

FROM DNA TO RNA

Transcription Produces RNA Complementary to One Strand of DNA

FROM DNA TO RNA

RNA Polymerases Carry Out Transcription

RNA Polymerases Carry Out Transcription

FROM DNA TO RNA

Cells Produce Different Categories of RNA Molecules

FROM DNA TO RNA

Signals Encoded in DNA Tell RNA Polymerase Where to Start and

Signals Encoded in DNA Tell RNA Polymerase Where to Start and

Signals Encoded in DNA Tell RNA Polymerase Where to Start and

Signals Encoded in DNA Tell RNA Polymerase Where to Start and

FROM DNA TO RNA

Transcription Start and Stop Signals Are Heterogeneous in

Transcription Start and Stop Signals Are Heterogeneous in

Transcription Start and Stop Signals Are Heterogeneous in

Transcription Start and Stop Signals Are Heterogeneous in

Transcription Start and Stop Signals Are Heterogeneous in

FROM DNA TO RNA

Transcription Initiation in Eukaryotes Requires Many Proteins

Transcription Initiation in Eukaryotes Requires Many Proteins

FROM DNA TO RNA

RNA Polymerase II Requires a Set of General Transcription Factors

Los factores de transcripcin generales

RNA Polymerase II Requires a Set of General Transcription Factors

RNA Polymerase II Requires a Set of General Transcription Factors

Secuencias consenso que se han encontrado cerca del inicio de la transcripcin

RNA Polymerase II Requires a Set of General Transcription Factors

FROM DNA TO RNA

La transcripcin por la RNA pol II en una clula eucarionte es ms compleja y

Polymerase II Also Requires Activator, Mediator, and ChromatinModifying Proteins

FROM DNA TO RNA

FROM DNA TO RNA

Transcription Creates Superhelical Tension

Transcription Creates Superhelical Tension

Transcription Creates Superhelical Tension

Transcription Creates Superhelical Tension

FROM DNA TO RNA

Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA

Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA

Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA

Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA

Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA

Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA

la transcripcin en eucariontes est acoplada al

El factor de transcripcin TFIIH fosforila el

Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA

Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA

FROM DNA TO RNA