From DNA to Protein: Genotype to Phenotype
From DNA to Protein: Genotype to Phenotype
• One Gene, One Polypeptide
• DNA, RNA, and the Flow of Information
• Transcription: DNA-Directed RNA Synthesis
• The Genetic Code
• Preparation for ________ Linking RNAs, Amino Acids, and Ribosomes
• Translation: RNA-Directed Polypeptide Synthesis
• Regulation of Translation
• Posttranslational Events
• Mutations: ________ Changes in Genes
One Gene, One Polypeptide
• A gene is defined as a DNA sequence.
• There are many steps between genotype and phenotype; genes cannot by themselves produce a phenotype.
One Gene, One Polypeptide
• In the 1940s, Beadle and Tatum showed that when an altered gene resulted in an altered phenotype, that altered phenotype always showed up as an altered enzyme.
• They experimented with strains of the bread mold Neurospora: a wild-type, and several mutant strains.
• Their results suggested that mutations cause a defect in only one enzyme in a metabolic pathway.
• This lead to the one-gene, one-enzyme hypothesis.


One Gene, One Polypeptide
• The gene–enzyme connection has undergone several modifications. Some enzymes are composed of different ________ coded for by separate genes.
• This suggests, instead of the one-gene, one enzyme hypothesis, a one-gene, one-polypeptide relationship.
DNA, RNA, and the Flow of Information
• The expression of a gene takes place in two steps:
ง ________ makes a single-stranded RNA copy of a segment of the DNA.
ง ________ uses information encoded in the RNA to make a polypeptide.
DNA, RNA, and the Flow of Information
• RNA (ribonucleic acid) differs from DNA in three ways:
ง RNA consists of only one polynucleotide strand.
ง The sugar in RNA is ________ not deoxyribose.
ง RNA has ________ instead of thymine.
• RNA can base-pair with single-stranded DNA (adenine pairs with uracil instead of thymine) and also can fold over and base-pair with itself.
DNA, RNA, and the Flow of Information
• Francis Crick's ________ ________ stated that DNA codes for RNA, and RNA codes for protein.
• How does information get from the nucleus to the cytoplasm?
• What is the relationship between a specific nucleotide sequence in DNA and a specific amino acid sequence in protein?

DNA, RNA, and the Flow of Information
• Messenger RNA, or mRNA moves from the nucleus of ________ cells into the cytoplasm, where it serves as a template for protein synthesis.
• Transfer RNA, or tRNA, is the link between the code of the mRNA and the amino acids of the polypeptide, specifying the correct amino acid sequence in a protein.

DNA, RNA, and the Flow of Information
• Certain viruses use RNA rather than DNA as their information molecule during transmission.
• These viruses transcribe from RNA to RNA; they make a complementary RNA strand and then use this "opposite" strand to make multiple copies of the viral genome by transcription.
• HIV and certain tumor viruses (called retroviruses) have RNA as their infectious information molecule; they convert it to a DNA copy inside the host cell and then use it to make more RNA.
Transcription: DNA-Directed RNA Synthesis
• In normal prokaryotic and eukaryotic cells, transcription requires the following:
ง A DNA template for ________ base pairing
ง The appropriate ribonucleoside triphosphates (ATP, GTP, CTP, and UTP) to act as substrates
ง The enzyme RNA polymerase
Transcription: DNA-Directed RNA Synthesis
• Just one DNA strand (the template strand) is used to make the RNA.
• For different genes in the same DNA molecule, the roles of these strands may be reversed.
• The DNA double helix partly unwinds to serve as template.
• As the RNA transcript forms, it peels away, allowing the already transcribed DNA to be rewound into the double helix.
Transcription: DNA-Directed RNA Synthesis
• The first step of transcription, ________ begins at a promoter, a special sequence of DNA.
• There is at least one promoter for each gene to be transcribed.
• The RNA polymerase binds to the promoter region when conditions allow.
• The promoter sequence directs the RNA polymerase as to which of the double strands is the template and in what direction the RNA polymerase should move.




Transcription: DNA-Directed RNA Synthesis
• After binding, RNA polymerase unwinds the DNA about 20 base pairs at a time and reads the template in the 3ข-to-5ข direction (elongation).
• The new RNA ________ from its 5ข end to its 3ข end; thus the RNA transcript is antiparallel to the DNA template strand.
• Transcription errors for RNA polymerases are high relative to DNA polymerases.

Transcription: DNA-Directed RNA Synthesis
• Particular base sequences in the DNA specify termination.
• Gene mechanisms for termination vary:
ง For some, the newly formed transcript simply falls away from the DNA template.
ง For other genes, a helper protein pulls the transcript away.
ง In prokaryotes, translation of the mRNA often begins before transcription is complete.

The Genetic Code
• A genetic code relates genes (DNA) to mRNA and mRNA to the amino acids of proteins.
• mRNA is read in ________ segments called codons.
• The number of different codons possible is 64 (43), because each position in the codon can be occupied by one of four different bases.
• The 64 possible codons code for only 20 amino acids and the start and stop signals.

The Genetic Code
• AUG, which codes for methionine, is called the start codon, the initiation signal for translation.
• Three codons (UAA, UAG, and UGA) are stop codons, which direct the ribosomes to end translation.
The Genetic Code
• After subtracting start and stop codons, the remaining 60 codons code for 19 different amino acids.
• This means that many amino acids have more than one codon. Thus the code is redundant.
• However, the code is not ambiguous. Each codon is assigned only one amino acid.
The Genetic Code
• In the early 1960s, molecular biologists broke the genetic code.
• Nirenberg prepared an artificial mRNA in which all bases were uracil (poly U).
• When incubated with additional components, the poly U mRNA led to synthesis of a polypeptide chain consisting only of phenylalanine amino acids.
• UUU appeared to be the codon for phenylalanine.
• Other codons were deciphered from this starting point.

Preparation for Translation:
Linking RNAs, Amino Acids, and Ribosomes
• The molecule tRNA is required to assure specificity in the translation of mRNA into proteins.
• The tRNAs must read mRNA correctly.
• The tRNAs must carry the correct amino acids.
Preparation for Translation:
Linking RNAs, Amino Acids, and Ribosomes
• The codon in mRNA and the amino acid in a protein are related by way of an adapter—a specific tRNA molecule.
• tRNA has three functions:
ง It carries an amino acid.
ง It associates with mRNA molecules.
ง It interacts with ribosomes.
Preparation for Translation:
Linking RNAs, Amino Acids, and Ribosomes
• A tRNA molecule has 75 to 80 nucleotides and a ________ shape (conformation).
• The shape is maintained by complementary base pairing and hydrogen bonding.
• The three-dimensional shape of the tRNAs allows them to combine with the binding sites of the ribosome.

Preparation for Translation:
Linking RNAs, Amino Acids, and Ribosomes
• At the 3ข end of every tRNA molecule is a site to which its specific ________ ________ binds covalently.
• Midpoint in the sequence are three bases called the anticodon.
• The anticodon is the contact point between the tRNA and the mRNA.
• The anticodon is complementary (and antiparallel) to the mRNA codon.
• The codon and anticodon unite by complementary base pairing.
Preparation for Translation:
Linking RNAs, Amino Acids, and Ribosomes
• Amino acids are attached to the correct tRNAs by activating enzymes called aminoacyl-tRNA synthetases.
• The enzyme has a three-part active site that binds:
ง A specific amino acid
ง ATP
ง A specific tRNA, charged with a high-energy bond
• The high-energy bond provides the energy for making the peptide bond.
Preparation for Translation:
Linking RNAs, Amino Acids, and Ribosomes
• The reactions have two steps:
ง Enzyme + ATP + AA ฎ enzyme—AMP—AA + PPi
ง Enzyme—AMP—AA + tRNA ฎ enzyme + AMP + tRNA—AA


Preparation for Translation:
Linking RNAs, Amino Acids, and Ribosomes
• Each ribosome has two subunits: a large one and a small one.
• In eukaryotes the large one has three different associated rRNA molecules and 45 different proteins.
• The small subunit has one rRNA and 33 different protein molecules.
• When they are not translating, the two subunits are separate.

Preparation for Translation:
Linking RNAs, Amino Acids, and Ribosomes
• The proteins and rRNAs are held together by ionic bonds and hydrophobic forces.
• The large subunit has four binding sites:
ง The T site where the tRNA first lands
ง The A site where the tRNA anticodon binds to the mRNA codon
ง The P site where the tRNA adds its amino acid to the polypeptide chain
ง The E site where the tRNA goes before leaving the ribosome
Preparation for Translation:
Linking RNAs, Amino Acids, and Ribosomes
• The small ribosomal subunit plays a role in validating the three-base-pair match between the mRNA and the tRNA.
• If hydrogen bonds have not formed between all three base pairs, the tRNA is ejected from the ribosome.
Translation: RNA-Directed Polypeptide Synthesis
• Translation begins with an initiation complex: a charged tRNA with its amino acid and a small subunit, both bound to the mRNA.
• This complex is bound to a region upstream of where the actual reading of the mRNA begins.
• The start codon (AUG) designates the first amino acid in all proteins.
• The large subunit then joins the complex.
• The process is directed by proteins called initiation factors.

Translation: RNA-Directed Polypeptide Synthesis
• Ribosomes move in the 5ข-to-3ข direction on the mRNA.
• The peptide forms in the N–to–C direction.
• The large subunit catalyzes two reactions:
ง Breaking the bond between the tRNA in the P site and its amino acid
ง Peptide bond formation between this amino acid and the one attached to the tRNA in the A site
• This is called peptidyl transferase activity.

Translation: RNA-Directed Polypeptide Synthesis
• After the first tRNA releases ________ it dissociates from the ribosome and returns to the cytosol.
• The second tRNA, now bearing a dipeptide, moves to the P site.
• The next charged tRNA enters the open A site.
• The peptide chain is then transferred to the P site.
• These steps are assisted by proteins called elongation factors.
Translation: RNA-Directed Polypeptide Synthesis
• When a stop codon—UAA, UAG, or UGA—enters the A site, a release factor and a water molecule enter the A site, instead of an amino acid.
• The newly completed protein then separates from the ribosome.

Regulation of Translation
• Antibiotics are defensive molecules produced by some fungi and bacteria, which often destroy other microbes.
• Some antibiotics work by blocking the synthesis of the bacterial cell walls, others by inhibiting protein synthesis at various points.
• Because of differences between prokaryotic and eukaryotic ribosomes, the human ribosomes are unaffected.
Regulation of Translation
• Polysomes are mRNA molecules with more than one ribosome attached.
• These make protein more rapidly, producing multiple copies of protein simultaneously.


Posttranslational Events
• Two posttranslational events can occur after the polypeptide has been synthesized:
ง The polypeptide may be moved to another location in the cell, or secreted.
ง The polypeptide may be modified by the addition of chemical groups, folding, or trimming.

Posttranslational Events
• As the polypeptide chain forms, it folds into its 3-D shape.
• The amino acid sequence also contains an "address label" indicating where in the cell the polypeptide belongs. It gives one of two sets of instructions:
ง Finish translation and be released to the cytoplasm.
ง Stall translation, go to the ER, and finish synthesis at the ER surface.
Posttranslational Events
• Polypeptides sent to the cytoplasm may contain information (signal sequences) that specifies a destination.
• The signal sequence binds to ________ proteins at the outer membrane of the appropriate organelle.
• A channel opens in the membrane, allowing the protein to pass through.
• In the process, the protein usually is unfolded by a ________ so that it can pass through the channel.
Posttranslational Events
• Polypeptides destined for the ER have a 25-amino-acid-long leader sequence.
• Before translation is finished, the leader sequence binds to a signal recognition particle.
• This stalls protein synthesis until the ribosome attaches to a specific receptor protein on the surface of the ER.
• Translation continues with the protein moving through a pore in the ER membrane.

Posttranslational Events
• Other signals are needed to direct further protein sorting:
ง Sequences of amino acids that allow the protein to stay in the ER
ง Sugars added in the Golgi apparatus to form glycoproteins, which go to lysosomes or the plasma membrane
• Proteins with no signals from the ER go through the Golgi apparatus and are secreted from the cell.
Posttranslational Events
• Most proteins are ________ after translation.
• These modifications are often essential to the functioning of the protein.
• Three types of modifications:
ง Proteolysis (cleaving)
ง Glycosylation (adding sugars)
ง Phosphorylation (adding phosphate groups)

Mutations: Heritable Changes in Genes
• Mutations are heritable changes in DNA—changes that are passed on to daughter cells.
• Multicellular organisms have two types of mutations:
ง Somatic mutations are passed on during mitosis, but not to subsequent generations.
ง Germ-line mutations are mutations that occur in cells that give rise to gametes.
Mutations: ________ Changes in Genes
• Some mutations, called conditional mutants, exert their effect only under certain restrictive conditions.
• A temperature-sensitive mutant allele, for example, may code for an enzyme that is altered at the restrictive temperature.
Mutations: Heritable Changes in Genes
• All mutations are alterations of the DNA nucleotide sequence and are of two types:
ง Point mutations are mutations of single genes.
ง Chromosomal mutations are changes in the arrangements of chromosomal DNA segments.
Mutations: Heritable Changes in Genes
• Point mutations result from the addition or subtraction of a base or the substitution of one base for another.
• Point mutations can occur as a result of mistakes during DNA replication or can be caused by environmental mutagens.
• Because of redundancy in the genetic code, some point mutations, called silent mutations, result in no change in the amino acids in the protein.

Mutations: Heritable Changes in Genes
• Some mutations, called missense mutations, cause an amino acid substitution.
• An example in humans is sickle-cell anemia, a defect in the b-globin subunits of hemoglobin.
• The b-globin in sickle-cell differs from the normal by only one amino acid.
• Missense mutations may reduce the functioning of a protein or disable it completely.


Mutations: Heritable Changes in Genes
• Nonsense mutations are base substitutions that substitute a stop codon.
• The shortened proteins are usually not functional.

Mutations: Heritable Changes in Genes
• A ________ ________ consists of the insertion or deletion of a single base in a gene.
• This type of mutation shifts the code, changing many of the codons to different codons.
• These shifts almost always lead to the production of nonfunctional proteins.

Mutations: Heritable Changes in Genes
• DNA molecules can break and re-form, causing four different types of mutations:
ง Deletions are a loss of a chromosomal segment.
ง Duplications are a repeat of a segment.
ง Inversions result from breaking and rejoining when segments get reattached in the opposite orientation.
ง Translocations result when a portion of one chromosome attaches to another.


Mutations: Heritable Changes in Genes
• Spontaneous mutations are permanent changes, caused by any of several mechanisms:
ง Nucleotides occasionally change their structure (called a tautomeric shift).
ง Bases may change because of a chemical reaction.
ง DNA polymerase sometimes makes errors in replication which can escape being repaired.
ง Meiosis is imperfect. Nondisjunction and translocations can occur.
Mutations: Heritable Changes in Genes
• Induced mutations are permanent changes caused by some outside agent (mutagen).
• Mutagens can alter DNA in several ways:
ง Altering covalent bonds in nucleotides
ง Adding groups to the bases
ง Radiation damages DNA:
q Ionizing radiation (X rays) produces free radicals.
q Ultraviolet radiation is absorbed by thymine and causes interbase covalent bonds to form.


Mutations: Heritable Changes in Genes
• Mutations have both benefits and costs.
• Germ line mutations provide genetic diversity for evolution, but usually produce an organism that does poorly in its environment.
• Somatic mutations do not affect offspring, but can cause cancer.
Mutations: Heritable Changes in Genes
• Mutations are rare events and most of them are point mutations involving one nucleotide.
• Different organisms vary in mutation frequency.
• Mutations can be detrimental, neutral, or occasionally beneficial.
• Random accumulation of mutations in the extra copies of genes can lead to the production of new useful proteins.

Video 12-01
Animation 12.1 Transcription
Animation 12.2 Deciphering the Genetic Code
Animation 12.3 Protein Synthesis