transcripción alberts 2014.pdf
TRANSCRIPT
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Molecular Biologyof the Cell
Sixth Edition
Chapter 6
How Cells Read the Genome:
From DNA to Protein
Copyright © Garland Science 2015
Alberts • Johnson • Lewis • Morgan • Raff • Roberts • Walter
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CHAPTER CONTENTS
FROM DNA TO RNA
FROM RNA TO PROTEIN
THE RNA WORLD AND THE ORIGINS OF LIFE
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Introduction
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Introduction
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FROM DNA TO RNA
• Introduction
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Introduction
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FROM DNA TO RNA
• RNA Molecules Are Single-Stranded
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RNA Molecules Are Single-Stranded
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RNA Molecules Are Single-Stranded
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RNA Molecules Are Single-Stranded
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RNA Molecules Are Single-Stranded
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RNA Molecules Are Single-Stranded
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RNA Molecules Are Single-Stranded
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RNA Molecules Are Single-Stranded
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RNA Molecules Are Single-Stranded
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RNA Molecules Are Single-Stranded
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FROM DNA TO RNA
• Transcription Produces RNA Complementary to One Strand of DNA
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Transcription Produces RNA Complementary to One Strand of DNA
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FROM DNA TO RNA
• RNA Polymerases Carry Out Transcription
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RNA Polymerases Carry Out Transcription
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RNA Polymerases Carry Out Transcription
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FROM DNA TO RNA
• Cells Produce Different Categories of RNA Molecules
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Cells Produce Different Categories of RNA Molecules
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FROM DNA TO RNA
• Signals Encoded in DNA Tell RNA Polymerase Where to Start and Stop
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Signals Encoded in DNA Tell RNA Polymerase Where to Start and Stop
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Signals Encoded in DNA Tell RNA Polymerase Where to Start and Stop
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Signals Encoded in DNA Tell RNA Polymerase Where to Start and Stop
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Signals Encoded in DNA Tell RNA Polymerase Where to Start and Stop
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FROM DNA TO RNA
• Transcription Start and Stop Signals Are Heterogeneous in Nucleotide Sequence
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Transcription Start and Stop Signals Are Heterogeneous in Nucleotide Sequence
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Transcription Start and Stop Signals Are Heterogeneous in Nucleotide Sequence
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Transcription Start and Stop Signals Are Heterogeneous in Nucleotide Sequence
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Transcription Start and Stop Signals Are Heterogeneous in Nucleotide Sequence
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Transcription Start and Stop Signals Are Heterogeneous in Nucleotide Sequence
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FROM DNA TO RNA
• Transcription Initiation in Eukaryotes Requires Many Proteins
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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.
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La RNA polimerasa II tiene muchas similitudes estructurales con la RNA polimerasa bacteriana, pero también
existen diferencias importantes:1- La RNA polimerasa bacteriana requiere solo de una proteína adicional para la iniciación de la transcripción,la RNA polimerasa II requiere de muchas proteínas adicionales,llamadas factores de transcripción generales.2- La iniciación de la transcripción en eucariotes trata con el empacamiento de DNA en nucleosomas yen cromatina, una estructura altamente organizada, ausente en el cromosoma bacteriano.
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Transcription Initiation in Eukaryotes Requires Many Proteins
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FROM DNA TO RNA
• RNA Polymerase II Requires a Set of General Transcription Factors
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RNA Polymerase II Requires a Set of General Transcription Factors
Los factores de transcripción generales
ayudan a posicionar a la RNA
polimerasa correctamente sobre el
promotor, ayudan a separar la doble
hélice para que comience la
transcripción y a liberar a la RNA
polimerasa del promotor al modo de
elongación, una vez que inició la
transcripción. Se dice que los factores
son generales porque se requieren en
casi todos los promotores de la RNApol
II.
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RNA Polymerase II Requires a Set of General Transcription Factors
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RNA Polymerase II Requires a Set of General Transcription Factors
Secuencias consenso que se han encontrado cerca del inicio de la transcripción
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RNA Polymerase II Requires a Set of General Transcription Factors
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FROM DNA TO RNA
• Polymerase II Also Requires Activator, Mediator, and Chromatin-Modifying Proteins
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• La transcripción por la RNA pol II en una célula eucarionte es más compleja y requiere muchas más proteínas.
• 1- requiere de proteínas regulatorias - activadores transcripcionales, los cuales deben unirse a secuencias de DNA específicas y ayudan a atraer la RNA pol II al punto de inicio de la transcripción
• 2- la iniciación de la transcripción en eucariontes in vivo requiere de un complejo protéico - mediador, que permite que las proteínas activadoras se comuniquen con la RNA pol II y con los factores de transcripción generales.
• 3- la iniciación de la transcripción 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.
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Polymerase II Also Requires Activator, Mediator, and Chromatin-Modifying Proteins
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FROM DNA TO RNA
• Transcription Elongation in Eukaryotes Requires Accessory Proteins
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FROM DNA TO RNA
• Transcription Creates Superhelical Tension
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Transcription Creates Superhelical Tension
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Transcription Creates Superhelical Tension
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Transcription Creates Superhelical Tension
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Transcription Creates Superhelical Tension
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FROM DNA TO RNA
• Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA Processing
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Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA Processing
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Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA Processing
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Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA Processing
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Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA Processing
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Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA Processing
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Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA Processing
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El factor de transcripción TFIIH fosforila el
CTD de la RNA polimerasa II, indicando la
transición de la iniciación a la elongación. Las
proteínas necesarias para el capping, para el
procesamiento y para la poliadenilación del pre
RNAm reconocen las diferentes formas
fosforiladas del CTD de la RNA polimerasa II.
De lo que se trata es que las proteínas
encargadas del capping, de procesamiento del
pre RNAm y de la poliadenilación del RNAm
se acerquen a sus sitios de acción en el
RNAm precursor conforme se transcriben
durante la elongación.
la transcripción en eucariontes está acoplada al
procesamiento del RNA
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Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA Processing
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Transcription Elongation in Eukaryotes Is Tightly Coupled to RNA Processing
Conforme la polimerasa transcribe, va cargando proteínas de procesamiento del preRNAm en su cola (CTD)las cuales va a transferir al RNAm naciente. La proteína de capping se une primero al CTD de la RNA polimerasa
cuando está fosforilada en la ser5 del hepta-repetido, una vez que la iniciación de la transcripción está avanzada. Esto asegura que la molécula 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 proteínas de elongación,y eventualmente se desfosforila en la ser5. Estas modificaciones en el CTD de la polimerasa van a atraer otras proteínasde procesamiento para que actúen 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 proteína crítica del procesamiento se ha transferido a la molécula de RNAm,
va a atraer al resto del complejo proteíco para que procese la molécula de RNA.
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FROM DNA TO RNA
• RNA Capping Is the First Modification of Eukaryotic Pre-mRNAs
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RNA Capping Is the First Modification of Eukaryotic Pre-mRNAs
3 enzimas, que actúan sucesivamente, realizan la reacción de capping.
Una fosfatasa elimina un fosfato del extremo 5’ nasciente del RNA,
una guanidiltransferasa añade 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 están listas para modificar el extremo 5’ naciente
al momento que emerge de la polimerasa.
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El cap 5’ metilado significa un extremo 5’ de un RNAm,
y esto le permite a la célula distinguir entre otros tipos de moléculas
de RNA presentes en la célula. La RNA polI y la RNA pol III no
producen transcritos con cap, en parte porque no tienen CTD.
Cual es la función del cap en el extremo 5’ del RNAm?
En el núcleo, el cap del 5’ del RNAm se une a un complejo protéico (CBC)
que le ayuda al mensajero a procesarse y exportarse adecuadamente.
También el cap tiene una función en la traducción.
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FROM DNA TO RNA
• RNA Splicing Removes Intron Sequences from Newly Transcribed Pre-mRNAs
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RNA Splicing Removes Intron Sequences from Newly Transcribed Pre-mRNAs
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RNA Splicing Removes Intron Sequences from Newly Transcribed Pre-mRNAs
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RNA Splicing Removes Intron Sequences from Newly Transcribed Pre-mRNAs
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RNA Splicing Removes Intron Sequences from Newly Transcribed Pre-mRNAs
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RNA Splicing Removes Intron Sequences from Newly Transcribed Pre-mRNAs
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RNA Splicing Removes Intron Sequences from Newly Transcribed Pre-mRNAs
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RNA Splicing Removes Intron Sequences from Newly Transcribed Pre-mRNAs
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FROM DNA TO RNA
• Nucleotide Sequences Signal Where Splicing Occurs
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Nucleotide Sequences Signal Where Splicing Occurs
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FROM DNA TO RNA
• RNA Splicing Is Performed by the Spliceosome
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El splicing de RNA está hecho por el spliceosoma
• Los pasos críticos en el procesamiento de RNA están hechos por
• moléculas de RNA- no por proteínas. Las moléculas de RNA reconocen las secuencias de nucleótidos que especifican los sitios de procesamiento y
también participan en la química del procesamiento (la catálisis).
• Son moléculas de RNA pequeñas (menos de 200nt) y 5 (U1, U2, U4, U5 y U6) son las que están involucradas en el tipo principal de splicing del pre-RNAm.
Son los snRNAs (RNAs pequeños nucleares), y cada uno forma un complejo con al menos 7 subunidades protéicas, y forman un snRNP (ribonucleoproteína
pequeña nuclear). Estas snRNPs forman la médula de spliceosoma, el complejo de RNA y proteínas que realiza el splicing.
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RNA Splicing Is Performed by the Spliceosome
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RNA Splicing Is Performed by the Spliceosome
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RNA Splicing Is Performed by the Spliceosome
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FROM DNA TO RNA
• The Spliceosome Uses ATP Hydrolysis to Produce a Complex Series of RNA–RNA Rearrangements
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The Spliceosome Uses ATP Hydrolysis to Produce a Complex Series of RNA–RNA Rearrangements
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FROM DNA TO RNA
• Other Properties of Pre-mRNA and Its Synthesis Help to Explain the Choice of Proper Splice Sites
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Other Properties of Pre-mRNA and Its Synthesis Help to Explain the Choice of Proper Splice Sites
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Other Properties of Pre-mRNA and Its Synthesis Help to Explain the Choice of Proper Splice Sites
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Other Properties of Pre-mRNA and Its Synthesis Help to Explain the Choice of Proper Splice Sites
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Other Properties of Pre-mRNA and Its Synthesis Help to Explain the Choice of Proper Splice Sites
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Other Properties of Pre-mRNA and Its Synthesis Help to Explain the Choice of Proper Splice Sites
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Other Properties of Pre-mRNA and Its Synthesis Help to Explain the Choice of Proper Splice Sites
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Other Properties of Pre-mRNA and Its Synthesis Help to Explain the Choice of Proper Splice Sites
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FROM DNA TO RNA
• Chromatin Structure Affects RNA Splicing
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FROM DNA TO RNA
• RNA Splicing Shows Remarkable Plasticity
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RNA Splicing Shows Remarkable Plasticity
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FROM DNA TO RNA
• Spliceosome-Catalyzed RNA Splicing Probably Evolved from Self-splicing Mechanisms
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FROM DNA TO RNA
• RNA-Processing Enzymes Generate the 3ʹ End of
Eukaryotic mRNAs
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RNA-Processing Enzymes Generate the 3ʹ End of Eukaryotic mRNAs
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RNA-Processing Enzymes Generate the 3ʹ End of Eukaryotic mRNAs
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RNA-Processing Enzymes Generate the 3ʹ End of Eukaryotic mRNAs
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RNA-Processing Enzymes Generate the 3ʹ End of Eukaryotic mRNAs
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FROM DNA TO RNA
• Mature Eukaryotic mRNAs Are Selectively Exported from the Nucleus
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Mature Eukaryotic mRNAs Are Selectively Exported from the Nucleus
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Mature Eukaryotic mRNAs Are Selectively Exported from the Nucleus
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Mature Eukaryotic mRNAs Are Selectively Exported from the Nucleus
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Mature Eukaryotic mRNAs Are Selectively Exported from the Nucleus
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Mature Eukaryotic mRNAs Are Selectively Exported from the Nucleus
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FROM DNA TO RNA
• Noncoding RNAs Are Also Synthesized and Processed in the Nucleus
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Noncoding RNAs Are Also Synthesized and Processed in the Nucleus
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Noncoding RNAs Are Also Synthesized and Processed in the Nucleus
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Noncoding RNAs Are Also Synthesized and Processed in the Nucleus
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Noncoding RNAs Are Also Synthesized and Processed in the Nucleus
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Noncoding RNAs Are Also Synthesized and Processed in the Nucleus
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FROM DNA TO RNA
• The Nucleolus Is a Ribosome-Producing Factory
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The Nucleolus Is a Ribosome-Producing Factory
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The Nucleolus Is a Ribosome-Producing Factory
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The Nucleolus Is a Ribosome-Producing Factory
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The Nucleolus Is a Ribosome-Producing Factory
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The Nucleolus Is a Ribosome-Producing Factory
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The Nucleolus Is a Ribosome-Producing Factory
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FROM DNA TO RNA
• The Nucleus Contains a Variety of Subnuclear Aggregates
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The Nucleus Contains a Variety of Subnuclear Aggregates
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The Nucleus Contains a Variety of Subnuclear Aggregates
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The Nucleus Contains a Variety of Subnuclear Aggregates
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The Nucleus Contains a Variety of Subnuclear Aggregates
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The Nucleus Contains a Variety of Subnuclear Aggregates
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FROM RNA TO PROTEIN
• An mRNA Sequence Is Decoded in Sets of Three Nucleotides
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An mRNA Sequence Is Decoded in Sets of Three Nucleotides
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An mRNA Sequence Is Decoded in Sets of Three Nucleotides
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FROM RNA TO PROTEIN
• tRNA Molecules Match Amino Acids to Codons in mRNA
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tRNA Molecules Match Amino Acids to Codons in mRNA
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tRNA Molecules Match Amino Acids to Codons in mRNA
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tRNA Molecules Match Amino Acids to Codons in mRNA
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tRNA Molecules Match Amino Acids to Codons in mRNA
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tRNA Molecules Match Amino Acids to Codons in mRNA
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tRNA Molecules Match Amino Acids to Codons in mRNA
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tRNA Molecules Match Amino Acids to Codons in mRNA
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FROM RNA TO PROTEIN
• tRNAs Are Covalently Modified Before They Exit from the Nucleus
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tRNAs Are Covalently Modified Before They Exit from the Nucleus
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tRNAs Are Covalently Modified Before They Exit from the Nucleus
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FROM RNA TO PROTEIN
• Specific Enzymes Couple Each Amino Acid to Its Appropriate tRNA Molecule
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Specific Enzymes Couple Each Amino Acid to Its Appropriate tRNA Molecule
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Specific Enzymes Couple Each Amino Acid to Its Appropriate tRNA Molecule
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Specific Enzymes Couple Each Amino Acid to Its Appropriate tRNA Molecule
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Specific Enzymes Couple Each Amino Acid to Its Appropriate tRNA Molecule
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Specific Enzymes Couple Each Amino Acid to Its Appropriate tRNA Molecule
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FROM RNA TO PROTEIN
• Editing by tRNA Synthetases Ensures Accuracy
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Editing by tRNA Synthetases Ensures Accuracy
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Editing by tRNA Synthetases Ensures Accuracy
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Editing by tRNA Synthetases Ensures Accuracy
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Editing by tRNA Synthetases Ensures Accuracy
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FROM RNA TO PROTEIN
• Amino Acids Are Added to the C-terminal End of a Growing Polypeptide Chain
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Amino Acids Are Added to the C-terminal End of a Growing Polypeptide Chain
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FROM RNA TO PROTEIN
• The RNA Message Is Decoded in Ribosomes
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The RNA Message Is Decoded in Ribosomes
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The RNA Message Is Decoded in Ribosomes
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The RNA Message Is Decoded in Ribosomes
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The RNA Message Is Decoded in Ribosomes
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The RNA Message Is Decoded in Ribosomes
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The RNA Message Is Decoded in Ribosomes
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The RNA Message Is Decoded in Ribosomes
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FROM RNA TO PROTEIN
• Elongation Factors Drive Translation Forward and Improve Its Accuracy
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Elongation Factors Drive Translation Forward and Improve Its Accuracy
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Elongation Factors Drive Translation Forward and Improve Its Accuracy
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Elongation Factors Drive Translation Forward and Improve Its Accuracy
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Elongation Factors Drive Translation Forward and Improve Its Accuracy
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FROM RNA TO PROTEIN
• Many Biological Processes Overcome the Inherent Limitations of Complementary Base-Pairing
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FROM RNA TO PROTEIN
• Accuracy in Translation Requires an Expenditure of Free Energy
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FROM RNA TO PROTEIN
• The Ribosome Is a Ribozyme
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The Ribosome Is a Ribozyme
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The Ribosome Is a Ribozyme
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The Ribosome Is a Ribozyme
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The Ribosome Is a Ribozyme
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The Ribosome Is a Ribozyme
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FROM RNA TO PROTEIN
• Nucleotide Sequences in mRNA Signal Where to Start Protein Synthesis
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Nucleotide Sequences in mRNA Signal Where to Start Protein Synthesis
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Nucleotide Sequences in mRNA Signal Where to Start Protein Synthesis
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Nucleotide Sequences in mRNA Signal Where to Start Protein Synthesis
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Nucleotide Sequences in mRNA Signal Where to Start Protein Synthesis
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FROM RNA TO PROTEIN
• Stop Codons Mark the End of Translation
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Stop Codons Mark the End of Translation
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FROM RNA TO PROTEIN
• Proteins Are Made on Polyribosomes
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Proteins Are Made on Polyribosomes
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Proteins Are Made on Polyribosomes
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Proteins Are Made on Polyribosomes
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FROM RNA TO PROTEIN
• There Are Minor Variations in the Standard Genetic Code
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There Are Minor Variations in the Standard Genetic Code
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FROM RNA TO PROTEIN
• Inhibitors of Prokaryotic Protein Synthesis Are Useful as Antibiotics
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Inhibitors of Prokaryotic Protein Synthesis Are Useful as Antibiotics
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Inhibitors of Prokaryotic Protein Synthesis Are Useful as Antibiotics
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FROM RNA TO PROTEIN
• Quality Control Mechanisms Act to Prevent Translation of Damaged mRNAs
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Quality Control Mechanisms Act to Prevent Translation of Damaged mRNAs
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FROM RNA TO PROTEIN
• Some Proteins Begin to Fold While Still Being Synthesized
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Some Proteins Begin to Fold While Still Being Synthesized
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Some Proteins Begin to Fold While Still Being Synthesized
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Some Proteins Begin to Fold While Still Being Synthesized
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FROM RNA TO PROTEIN
• Molecular Chaperones Help Guide the Folding of Most Proteins
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FROM RNA TO PROTEIN
• Cells Utilize Several Types of Chaperones
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Cells Utilize Several Types of Chaperones
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Cells Utilize Several Types of Chaperones
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Cells Utilize Several Types of Chaperones
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Cells Utilize Several Types of Chaperones
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FROM RNA TO PROTEIN
• Exposed Hydrophobic Regions Provide Critical Signals for Protein Quality Control
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Exposed Hydrophobic Regions Provide Critical Signals for Protein Quality Control
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FROM RNA TO PROTEIN
• The Proteasome Is a Compartmentalized Protease with Sequestered Active Sites
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The Proteasome Is a Compartmentalized Protease with Sequestered Active Sites
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The Proteasome Is a Compartmentalized Protease with Sequestered Active Sites
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The Proteasome Is a Compartmentalized Protease with Sequestered Active Sites
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The Proteasome Is a Compartmentalized Protease with Sequestered Active Sites
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The Proteasome Is a Compartmentalized Protease with Sequestered Active Sites
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The Proteasome Is a Compartmentalized Protease with Sequestered Active Sites
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The Proteasome Is a Compartmentalized Protease with Sequestered Active Sites
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FROM RNA TO PROTEIN
• Many Proteins Are Controlled by Regulated Destruction
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Many Proteins Are Controlled by Regulated Destruction
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Many Proteins Are Controlled by Regulated Destruction
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Many Proteins Are Controlled by Regulated Destruction
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FROM RNA TO PROTEIN
• There Are Many Steps From DNA to Protein
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There Are Many Steps From DNA to Protein
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THE RNA WORLD AND THEORIGINS OF LIFE
• Introduction
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Introduction
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THE RNA WORLD AND THEORIGINS OF LIFE
• Single-Stranded RNA Molecules Can Fold into Highly Elaborate Structures
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Single-Stranded RNA Molecules Can Fold into Highly Elaborate Structures
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Single-Stranded RNA Molecules Can Fold into Highly Elaborate Structures
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Single-Stranded RNA Molecules Can Fold into Highly Elaborate Structures
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Single-Stranded RNA Molecules Can Fold into Highly Elaborate Structures
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Single-Stranded RNA Molecules Can Fold into Highly Elaborate Structures
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Single-Stranded RNA Molecules Can Fold into Highly Elaborate Structures
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THE RNA WORLD AND THEORIGINS OF LIFE
• RNA Can Both Store Information and Catalyze Chemical Reactions
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RNA Can Both Store Information and Catalyze Chemical Reactions
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THE RNA WORLD AND THEORIGINS OF LIFE
• How Did Protein Synthesis Evolve?
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THE RNA WORLD AND THEORIGINS OF LIFE
• All Present-Day Cells Use DNA as Their Hereditary Material
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All Present-Day Cells Use DNA as Their Hereditary Material
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PROBLEMS
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PROBLEMS
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PROBLEMS
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PROBLEMS
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PROBLEMS
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PROBLEMS