Today is Thursday,January 8th, 2015

Pre-Class:DNA makes…what?

In This Lesson:Protein Synthesis

(Lesson 2 of 3)

Today’s Agenda

• The Central Dogma• Transcription• Translation• Mutations• Arts and Crafts• The Longest Word Ever

• Where is this in my book?– Chapters 17-18.– NOTE: This lesson can get very complicated.– USE YOUR TEXTBOOK AND ESPECIALLY THE GLOSSARY.

By the end of this lesson…

• You should be able to describe in full detail how a DNA gene becomes a protein and thus a trait.

• You should be able to explain how those genes are turned on and off.

• You should be able to analyze the effects of a mutation on the genetic code.

The Big Idea• Remember those experiments to determine whether

DNA or proteins were the basics of heredity?– Damn skippy you do.

• Well, as you know, there’s a very close relationship between proteins and DNA.

• So close, in fact, that it is perhaps even more of a “core idea” to biology than is evolution.– We call it the central dogma (of biology).

• The word “dogma” is usually used to describe the underlying principles of a religion that are never doubted. That should tell you something about how big a deal this is.

The Central Dogma

NOTE: DNA Replication does not have to happen before transcription.

DNA TraitProteinRNA

Transcription Translation

Replication

The Central Dogma

DNA TraitProteinRNA

Transcription Translation

Nucleus(if eukaryotic)

Cytoplasm @ Ribosome

Big Ideas from the Big Idea

• From the central dogma it should be obvious that:– This is the mechanism for DNA to endow an

organism with traits.– The nucleus speaks in DNA, the ribosome speaks

in RNA.– DNA gets a whole lot of credit but it’s protein that

does all the work.

Where We’re Going

• Following the central dogma, we’re going to review how we get from DNA to RNA to protein.

• Then we’ll look at what happens when something goes wrong (as usual).

The Origin of Traits

• So you can see that strong early evidence for the effect of genes and role of proteins was provided by metabolism.

• If the DNA responsible for production of particular enzymes is non-functional, the damaged metabolic pathway will reveal itself with a particular phenotype.

• Here’s what I mean…

Sample Metabolic PathwayIngested Protein

Digestion

Phenylalanine

Phenylalanine hydroxylase

Tyrosine

Transaminase

Hydroxyphenylpyruvic acid

Hydroxyphenylpyruvic acid oxidase

Homogentisic acid

✖

Phenylketonuria

✖

Tyrosinosis

Melanin

Thyroxine

Cretinism

✖

Albinism✖

The Origin of Traits

• All of this phenotypic analysis led researchers to develop the one gene, one enzyme hypothesis.– Damaged DNA will express itself in the form of a

nutritional disorder, allowing the gene to be mapped.

– Of course, this isn’t entirely true for all genes, but it helped in the early days.

The Process

• Okay, let’s start reviewing the process of transcription.

• Transcription is the creation of mRNA from DNA.

• RNA (including mRNA) has three main differences from DNA.

• Do you remember them?

About RNA…

• RNA is another kind of nucleic acid.– Ribonucleic acid.

• It is usually single-stranded, DNA is double-stranded.

• Instead of having Deoxyribose, it has Ribose.

• Instead of having Thymine, it has Uracil.

Mmm, sugar!

• Sugar differences:– Deoxyribose is missing an

oxygen.

http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/P/Pentose.gif

Compare and Contrast• Using these pictures and your notes, organize

a T-chart comparing DNA and RNA.

Here’s what I got…

DNA RNA

Bases A, C, G, T A, C, G, U

Sugar Deoxyribose Ribose

Strands Two One

Types of RNA

• There are several types of RNA out there as well, with the main three being:– mRNA (messenger RNA)– tRNA (transfer RNA)– rRNA (ribosomal RNA)

• Remember these and watch for them to arise later throughout transcription and translation.

• FYI, all three are made from DNA.

The Process of Transcription

• Just like in replication, the DNA is unwound, but this time by RNA polymerase.

• RNA nucleotides begin to be added to a new mRNA strand.– This time, not by DNA polymerase but…– RNA polymerase again! (the enzyme that does it all)

• Key: Every time the mRNA molecule would have a T, a U for Uracil is placed instead.– Uracil is also a pyrimidine.

Transcription---H---

---H---

---H---Deoxy-ribose

P

Thymine

Deoxy-ribose

P

Cytosine

Deoxy-ribose

Adenine

RNA Polymerase breaks H-bondsStrands move apartRNA Polymerase makes mRNAmRNA exits nucleusDNA re-coils

Deoxy-ribose

P

Thymine

Deoxy-ribose

P

Guanine

Deoxy-ribose

P

AdenineRibose

Adenine

P

RiboseGuanine

P

RiboseUracil

P

RiboseAdenine

P

RiboseGuanine

P

RiboseUracil

P

---H---

---H---

---H---

Transcription Details

• Instead of a replication bubble, RNA polymerase opens a transcription bubble.

• RNA is polymerized 5’ to 3’, just like DNA.• The strand of DNA that is used for transcription is

the antisense strand (or template strand or reading strand).

• The strand of DNA not used for transcription is the sense strand (or coding strand).– It has the same sequence as the mRNA strand.

---H---

---H---

---H---Deoxy-ribose

P

Thymine

Deoxy-ribose

P

Cytosine

Deoxy-ribose

Adenine

Thymine

Guanine

AdenineRibose

Adenine

P

RiboseGuanine

P

RiboseUracil

P

RiboseAdenine

P

RiboseGuanine

P

RiboseUracil

P

---H---

---H---

---H---

Sense Strand Antisense Strand

Practice

• Gene Expression – Transcription POGIL– Up to first STOP sign – we’re going to go over this

one!

The Road Ahead

• It’s about to get complicated up in hurr…• What I’m going to do is give you some key

vocabulary words now, then repeat them exactly later on.

• You can write them now, later, or in context as I reach the concepts during the presentation.

One other thing…

• Much of the focus of this lesson, more so than when you covered this in previous biology classes, is concerned with control.

• How does a cell control transcription?• How do we prevent, say, digestive stomach

enzymes from being manufactured in the brain?– Your brain cells have those genes, after all.– Every cell has all the genes.

Vocabulary Summary[same as later]

• Promoter– DNA region where RNA polymerase attaches and transcription starts.

• Operator– Prokaryotic DNA region to which a repressor may bind.

• Repressor– A protein that attaches to DNA and blocks transcription.

• Prokaryotes: Near or in promoter.• Eukaryotes: Could be near the promoter, could be far away.

• Activator– A protein that attaches to DNA and aids transcription.

• Prokaryotes: Near or in promoter.• Eukaryotes: Could be near the promoter, could be far away.

• Terminator– Prokaryotic DNA region that signals transcription to stop.

• Closest eukaryotic equivalent is the polyadenylation signal.

Transcription Details

• Like DNA replication, we know a bit more about transcription from studying prokaryotes, but it’s similar in eukaryotes too.

• Here’s the sequence:1. Initiation• There’s going to be a lot on this one.

2. Elongation3. Termination

1. Initiation: Prokaryotes

• Meet Barry the Bacterium.– He’s E. coli, just so we’re

clear.• He likes lactose.– The sugar found in milk.

• He doesn’t get it all the time, though, and he needs a special group of enzymes to digest it.

• Barry’s got a “choice” to make.

Mmmmm!I lurrves me some lactose!

Barry’s Choice

• Help a bacterium out. Barry can do one of two things:– Barry can make those lactose-related enzymes all the

time, and then use them every once in a while when he gets some lactose.

– Barry can wait till he gets lactose, then make some of those enzymes.

• Which makes more sense biologically?– To put it another way, do you get all the groceries you

need for the year or get them on an as-needed basis periodically?

Barry Made the Right Choice

• Transcription and translation cost energy, so it’s best to do them only when necessary.

• Prokaryotes use something called a repressor to block several genes for lactose-related enzymes, which we collectively call the lac operon.

If I were a real bacterium,I wouldn’t have a face.

Simulation

• PhET: Gene Machine – lac Operon

About Operons• Operons are groups of genes controlled by a single

region of DNA called a promoter.– The promoter is simply an area upstream of the genes

where RNA polymerase can bind to the DNA.• In between the promoter and the genes is the

operator.– The operator is a stretch of DNA responsible for allowing or

denying transcription. How?

lac z lac y lac a

Lactose-Related Genes

OperatorPromoter

About Operons• Attached to the operator may be a repressor, which is a

protein molecule that simply blocks RNA polymerase.• Here’s the cool part:– For the lac operon, the presence of lactose removes the repressor.– RNA polymerase is then able to transcribe the relevant genes,

which includes an enzyme that degrades lactose.• Thus, this is negative feedback.• Lactose removes the repressor so RNA polymerase makes lactase, which

removes lactose.

lac z lac y lac a

Lactose-Related Genes

OperatorPromoter

RepressorRNA Polymerase LactoseRNA Polymerase

Removing the Repressor

Repressor leaving mRNA.http://www.ncbi.nlm.nih.gov/bookshelf/br.fcgi?book=cooper&part=A964&rendertype=figure&id=A974

Other Uses of Operons• In the previous case, the presence of lactose stimulated a gene

to be expressed.• Sometimes the absence of a substance will start gene

expression.• For example, the trp operon controls five genes that make

tryptophan, an amino acid.• The repressor detaches when tryptophan is not present.

– This is also an example of negative feedback.

trp e trp d trp c

Tryptophan-Producing Genes

OperatorPromoter

Repressor

trp b trp a

TryptophanRNA Polymerase

Trp Operon Metaphor

• Think of the trp operon like a generator you’d hook up to a house or to a building.

• You don’t want it running most of the time, unless you’re out of power.– Then you want it on.

• In the same way, the trp operon is off when the cell is getting tryptophan, but when tryptophan runs out, the operon is activated.– Once the cell makes enough tryptophan, the amino acid

(tryptophan) itself will reactivate the repressor and turn the operon off again.

Operon Regulation

• Inducible operons are those that are turned on when an inducer is present.– For the lac operon, its repressor is deactivated by the

presence of allolactose (the inducer).• Allolactose’s presence deactivates the repressor.

• Repressible operons are those that are turned off when something is present.– For the trp operon, tryptophan’s presence activates

the repressor.

Positive versus Negative Control

• Both the trp and lac operons are examples of negative control.– Why negative?– Because an active repressor switches them off.

• Positive control is when the presence of something activates an activator.– An activator is something that will promote the

attachment of RNA polymerase.– An activator turns the genes on.

Initiation in Eukaryotes

• Eukaryotes are similar, but may sometimes use an activator and transcription factors that need to bind to get things started.– We still use repressors, too. Sometimes.– We do not use operons, or at least they haven’t

been discovered in us yet.• Instead, we use little complexes that together are called

transcription units. They’re similar to operons.

1. Initiation: Eukaryotes

• A bunch of transcription factors bind to a promoter upstream of the actual gene and signal RNA polymerase II to bind.– Transcription factors are proteins that

turn transcription on and off.– How do they know where to bond?

The promoter region has in it a TATA box.• The TATA box is actually a stretch of

“TATA” followed by “A.”

• Combined, the RNA polymerase and transcription factors make up the transcription initiation complex.

Video!

• Transcription – Advanced

Further Eukaryotic Differences• Okay, here’s where it gets a little

complicated.– Note that these next few slides will not

be covered on the test.• Eukaryotes have two major

“curveballs” thrown into the transcription process:– The most obvious one is the nucleus.– The second one is the structure of

chromatin.• Remember that chromatin is DNA

wrapped around spherical histone proteins.

• The histone/DNA complex is called a nucleosome.

https://www.broadinstitute.org/files/news/images/2010/chromatin_states_2a.png

Chromatin video

Eukaryotic Initiation

• The first thing that needs to happen is that the histones need to get out of the way.– Yep, enzymes do it.– Histone acetylation (adding acetyl groups) opens the

DNA for transcription.– Histone methylation (adding –CH3 methyl groups)

closes the DNA and makes transcription less likely.• Remember from way back in our organic chemistry stuff –

methyl groups tend to inactivate stuff.

Eukaryotic Initiation

• Sometimes genes themselves are methylated.– The “letters” are not changed, but they’re “tagged”

with –CH3 groups.– This methylation stays with the genes and can be

passed on to offspring.– This is the foundation of epigenetics (inheritance via

means other than the nucleotide sequence of DNA).– Epigenetic inheritance is reversible and explains

genomic imprinting.• Remember that? When a set of one parent’s genes are

shut down?

Back to Initiation

• Eukaryotic DNA has a lot of control elements, which are sections of DNA to which transcription factors bind.– Transcription factors, not RNA polymerase itself.

• These control elements can be all over the place in the DNA strand.– Proximal control elements are near the gene promoter.– Distal control elements are far from the gene promoter.

• Groups of distal control elements are called enhancers.

• Key: All of these components need to come together to start transcription.

So what’s the difference?

• Prokaryotes’ operons tend to be relatively simple.

• Eukaryotes’ genes may be regulated by myriad combinations of control elements and transcription factors, allowing for all kinds of different gene expression.– Yep, genetics is complicated and awesome.

Eukaryotic Initiation

http://www.nature.com/nrm/journal/v5/n5/images/nrm1369-i1.gif

Initiation Summary

• Prokaryotes– RNA polymerase attaches to a promoter and

transcribes genes.– It may be stopped by a repressor attached to the

operon.• Eukaryotes– Transcription factors (proteins) bind to the promoter.• The promoter has a region in it called the TATA box.

– Once the transcription factors have bonded, then RNA polymerase II can bind.

2. Elongation

• RNA polymerase transcribes DNA into mRNA as it unwinds.– Speed is ~20 bases a second.

• Typical genes are 300-500 bases total.

– Error rate is ~1 in 100,000 bases.• However, since this is mRNA

and not a more permanent DNA, error checking generally isn’t worth it.

3. Termination

• RNA polymerase stops at a termination sequence.– This is different for prokaryotes and eukaryotes – more on

this on the next slide.

RNA GC hairpin turn

3. Termination Eukaryote Difference

• In prokaryotes, RNA polymerase releases at the termination signal (called the terminator).

• In eukaryotes, RNA polymerase II eventually reaches a polyadenylation signal.– The signal is “AAUAAA.”

• It then sails past the signal for hundreds of nucleotides.– The actual mRNA (not RNA polymerase II) is cut free

after about 10 to 35, however.– RNA polymerase II makes more mRNA that just gets

degraded until enzymes catch up to it and release it.

Further Differences

• In eukaryotes, transcription is accomplished using three different RNA polymerase enzymes:– RNA Polymerase I makes rRNA.

• More on that later.

– RNA Polymerase II makes mRNA.• What we just learned.

– RNA Polymerase III makes tRNA.• More on that later.

• They function differently by only responding to certain promoter sequences in the DNA.

Vocabulary Summary[same as before]

• Promoter– DNA region where RNA polymerase attaches and transcription starts.

• Operator– Prokaryotic DNA region to which a repressor may bind.

• Repressor– A protein that attaches to DNA and blocks transcription.

• Prokaryotes: Near or in promoter.• Eukaryotes: Could be near the promoter, could be far away.

• Activator– A protein that attaches to DNA and aids transcription.

• Prokaryotes: Near or in promoter.• Eukaryotes: Could be near the promoter, could be far away.

• Terminator– Prokaryotic DNA region that signals transcription to stop.

• Closest eukaryotic equivalent is the polyadenylation signal.

Okay, time for a “kindabreak.”

• Here’s what you need to do:– Get a piece of masking tape long enough to write the

words “APproPRIAteLY JOinED” on it.– Write the words “APproPRIAteLY JOinED” on it exactly

like that, leaving slightly more space between the letters than you normally would.

– Now tear the piece of masking tape between letters of different cases (i.e. between “o” and “P”).

– Tape only the lower case letters to your desk in sequence.

– What did you spell?

mRNA Processing

• In prokaryotes, all of the mRNA that is transcribed is used, but in eukaryotes, there are stretches of DNA that are transcribed into mRNA but are later removed.– Introns are the parts of the mRNA removed.• These parts interrupt (like your CAPITAL letters).

– Exons are the parts of the mRNA that are kept.• These parts are expressed (like your lower case letters).

Why introns?

• So why make introns at all if they’re just going to be removed?

• It turns out that different combinations of introns and exons, even from the same stretch of DNA, can ultimately make different proteins.– This is called alternative RNA splicing.– Case in point: As you’ll see in your POGIL, humans have

25,000 genes but make 100,000 proteins.– Introns/exon combinations explain the variability.

mRNA Processing

• The original, complete mRNA molecule is called the primary mRNA transcript (or pre-mRNA):

• Then the introns are excised (confusing?) using spliceosomes.– Spliceosomes are proteins composed of small nuclear

ribonucleoproteins, or snRNPs (pronounced “snurps”).• Once edited, the mRNA is known as the mature

mRNA transcript:

• But what are those things on the ends?

Further Processing• A 5’ cap and a poly-A tail are then added.– The 5’ cap is a GTP molecule for protection/signaling.– The poly-A tail is a stretch of 50 to 250 “A” bases for protection and

aid in transport out of the nucleus.• The poly-A tail is added by poly A polymerase.• Remember the polyadenylation signal?

Further Processing• In order, a completed eukaryotic mRNA molecule has:

– 5’ Cap– Leader (space between cap and start codon)– Start Codon (AUG – methionine – more later)– Stop Codon (UAA/UAG/UGA – more later)– Trailer (space between stop codon and poly-A tail)

• The polyadenylation signal sequence is in the trailer.

– Poly-A Tail• Note: Technically, this mRNA processing happens virtually

instantaneously as mRNA is transcribed.

mRNA Processing

https://www.mpibpc.mpg.de/2224704/Fig1_eng_jpg.jpg

One last thing on RNA splicing…

• It appears that the sequence of exons in a protein leads to different domains once the peptide is completed.– Domains are different

functional sections of the protein.

Whew.

• That’s a lot. Let’s recap:– Spliceosomes animation

Stop Codons, Trailers, and Terminators

• In case you’re wondering:– A stop codon tells the ribosome to stop translating –

that’s not a part of transcription.– A trailer is a region of “concluding mRNA.”• It’s not made into proteins.

– A termination signal ends production of the intended mRNA strand.• In eukaryotes, RNA polymerase II keeps going and makes a

new strand that has no 5’ cap so it gets degraded.• Eventually, enzymes catch up to RNA polymerase and

detach it from the DNA.

Sidney AltmanYale University

Thomas CechUniversity of

Colorado

Aside: RNA Self-Splicing?• Remember when we talked

about ribozymes, those little bits of RNA that can act as enzymes?

• Well, they’re capable of self-splicing, suggesting that they act as an enzyme.

• As we learned before, ribozyme RNA may be one of the earliest adaptations of life.– This stuff was discovered in the

1980s by Sidney Altman and Thomas Cech.

What happens to the mRNA strand?

• In case you’re wondering, the mRNA strand, upon exiting the nucleus, is…

• …IMMEDIATELY IN PERIL!• Enzymes called exonucleases break down the

mRNA strand starting at the 3’ end.– That poly-A tail makes a lot of sense now, right?– The poly-A tail prevents the “good” parts of the mRNA

strand from being immediately degraded.• The free RNA bases can be used again during

another round of transcription.

Practice

• Gene Expression – Transcription POGIL– Finish it – we’re going to go over it!

Now then…

• …let’s get to the actual protein synthesis.• Up until this point we’ve talked about the process

of transcription, or processes associated with transcription like RNA splicing.– Remember, transcription is DNA mRNA.

• Now let’s cover the next step: mRNA Protein.• And which organelle is responsible for protein

synthesis?– That’s right, ribosomes.– Good job [insert student name here].

Translation

• Going from mRNA to protein is the process of translation.

• In prokaryotes, since there is no RNA processing, transcription and translation happen simultaneously.

Transcription

Translation

mRNA

Protein

Bacterial Chromosome

Translation in Prokaryotes

• As the mRNA strand grows, ribosomes attach to it and create protein.– By the way, a

group of ribosomes reading mRNA concurrently is called a polyribosome.

Translation in Eukaryotes

• In eukaryotes, the process takes a bit longer due to two main “separations:”– Time• mRNA must be processed.• It takes about an hour to go from

DNA to protein.

– Location• Eukaryotes have nuclei; the mRNA

must exit the nucleus before translation can begin.

Translation in Eukaryotes

• Here’s a nice summary image. Look for things that are familiar.

Translation: The General Process

• Translation can be summarized in two general steps.

• Obviously, these each have their own details, but it really comes down to this:1. mRNA exits the nucleus through a nuclear pore.2. Two halves of a ribosome assemble on the mRNA

and use its “instructions” to make protein.• Some quick reminders:– Protein monomer = amino acid.– Protein polymer = polypeptide.

Universal Genetic Code

• Here’s where you need a reference table.• Universal Genetic Code [found in Fact Sheets]

Codons

• Every three mRNA bases read by the ribosome is called a codon.

• Each codon “codes for” a particular amino acid.– Except for the three stop codons.

• Example: AUG CCC GAU UCG UGA– AUG = Methionine (always the start)– CCC = Proline– GAU = Aspartate– UCG = Serine– UGA = [STOP]

Translation Example

• Here’s my mRNA sequence:– CGGAUGCUGAAGUGAGGC

• Now here it is with the codons separated:– CGG | AUG | CUG | AAG | UGA | GGC

• Now here it is when the amino acids linked together:– Methionine-Leucine-Lysine

• Notice how I did not start at the beginning (not a start codon) and stopped at the stop codon.

Aside: The Discovery of Codons

• Francis Crick (that Crick) determined the triplet codon concept.

• In the 1960s, Marshall Nirenberg and Har Gobind Khorana figured out that codons were linked to particular amino acids.– They made a synthetic mRNA strand

of entirely U’s.– UUU was then determined to code

for phenylalanine.http://assets.upstart.bizjournals.com/images/site/editorial/magazine/23andme-timeline-02-slide*580.jpg?v=1

Khorana Nirenberg

Codons

• Take a look at your universal genetic code sheet.• Here are some things to note:– Because there are 4 different bases and 3 letters in a

codon, there are 64 different possible codons.• 4 * 4 * 4 = 64.

– The code is redundant (sometimes called degenerate).• Most amino acids are linked to multiple codons.

– For many amino acids, only the first two letters are important.• Take a look at the letter combinations that make proline, for

example. Once you get the first two letters, the last doesn’t matter.

• We’ll look further into this concept when we explore “wobble.”

Practice

• Gene Expression – Translation POGIL– Up to first STOP sign – we’re going to go over it!

Translation• So how do codons code for amino acids?– With another form of RNA called tRNA (transfer RNA).

• tRNA averages around 76 nucleotides of RNA in length.

– Transfer RNA has a cloverleaf structure and has two especially important parts:• An amino acid attachment site.• A three-base anticodon.

Anticodons

• As you might have guessed, the anticodon on tRNA is the opposite of the codon on mRNA.– Example codon: AUG– Corresponding anticodon: UAC

• Thus, tRNA appears at the correct codon because it has, to use the old expression, the other half of the BFF necklace.– tRNA brings to the growing polypeptide the

correct amino acid.

Wobble

• A moment ago I mentioned how there are (in some cases) many codons for the same amino acid.

• Part of the reason why is that the third base in each codon may not be entirely necessary.

• Crick (yep, him) discovered that the third base doesn’t have the same base-pairing “strictness” as the other two bases do.

Wobble

• Furthermore, tRNA actually has other bases besides A, U, C, G that sometimes act like wild cards.– Mainly I, for inosine.• I serves as a “match” for A, U, or G.

• The third base is therefore “wobbly” and isn’t quite so essential.– It’s actually called “the wobble base.”

Wobble

http://www.soc-bdr.org/rds/authors/unit_tables_conversions_and_genetic_dictionaries/e5202/index_en.html

First or Second Letter in mRNA(5’ to 3’)

First or Second Letter in tRNA(3’ to 5’)

A UU AC GG C

Third Letter in mRNA (5’ to 3’) Third Letter in tRNA (3’ to 5’)A or G U

U AU or C G

G CU, C, A I

Last thing on wobble…

• In case you need more evidence for wobble, keep in mind that there are 61 different mRNA codons (excluding “stop”), yet 45 different tRNA molecules.

• Explain that one without using wobble…

Replication, Transcription, Translation

DNAAntisense Strand

DNASense Strand

mRNACodon

tRNAAnticodon Amino Acid

TTG AAC AAC UUG Asparagine

AGT TCA UCA AGU Serine

TCG AGC AGC UCG Serine

ACA TGT UGU ACA Cysteine

ACC TGG UGG ACC Tryptophan

tRNA• tRNA is “loaded” with amino acids by an enzyme called

aminoacyl tRNA synthetase.– ATP is hydrolyzed to AMP and the energy from those bonds is

stored in the tRNA-amino acid bond.– It’s unstable and high-energy, so the amino acid can “pop off”

easily.• That’ll be important later.

Aminoacyl tRNA Synthetase

• Notice how two phosphate groups are used to “load” a tRNA molecule with an amino acid.

Summary

• Okay, let’s do a little recap.• DNA is used to make mRNA in transcription.• mRNA is processed.– Addition of 5’ cap.– Addition of poly-A tail.– Removal of introns.

• mRNA exits the nucleus.• tRNA matches every three bases on the mRNA (a

codon) with its own anticodon (three tRNA bases) and brings over the appropriate amino acid.

Coordination

• But wait…where is all this happening? How is this process being coordinated?

• Key: The process of translation is governed by the ribosome.– Ribosomes are made of a third kind of RNA called rRNA.– They have two parts – a large and a small subunit.– They also have three “sites” used in reading the mRNA.• Let’s look closer…

Aside: Ribosome Differences

• Turns out that ribosomes are slightly different in prokaryotes and eukaryotes.

• Widely-used antibiotics Streptomycin and Tetracyclin work by disabling prokaryotic ribosomes.

Ribosome Sites• A Site– Aminoacyl-tRNA site– tRNA in this site has the amino acid waiting to be added to the

polypeptide chain.• Memory Device: The A site has the awaiting amino acid as it arrives.

• P Site– Peptidyl-tRNA site– tRNA in this site has the polypeptide chain.

• Memory Device: The P site has the polypeptide as it polymerizes.

• E Site– Exit site– tRNA in this site does not have an amino acid and will leave the

ribosome.

Large Ribosomal Subunit (rRNA)E Site A SiteP Site

Small Ribosomal Subunit (rRNA)

tRNA

U A C

Translation Mechanism

MET

tRNA

U A U

A U G A C CA UA G C A U G A AA UC

ISO

MET

A U G A C CA UA G C A U G A AA UC

tRNA

U A U

tRNA

U A C

tRNA

G G G

PRO

mRNA

This process continues until a

stop codon is reached, at which point the mRNA

strand, tRNA units, and rRNA subunits

all are released.Start Codon (Methionine)

Translation Sequence• The process of translation is divided into

three steps similar to transcription. (leave room in your notes for more!)1. Initiation

• mRNA, tRNA, rRNA come together.

2. Elongation• Amino acids are added as the polypeptide grows.

3. Termination• The stop codon is reached.

1. Initiation Details• Initiation factors are small proteins that bring

together tRNA and the small rRNA subunit at the start codon of mRNA, methionine.

• The large subunit arrives just afterward.

2. Elongation Details

• Elongation factors bring the correct tRNA into the A site.

• rRNA catalyzes the formation of a peptide bond between the two amino acids (A site and P site).– GTP is used for power.

• The ribosome moves down the mRNA and the third tRNA exits from the E site.

3. Termination • A protein known as a release factor binds to codons UAA, UAG,

and UGA – the stop codons.• Water is added to the end of the polypeptide chain to end it.• Proteins may then undergo post-translational modification

before they are functional.

A reminder…

• Remember what the rough endoplasmic reticulum does?– The rough ER makes proteins…for export from the

cell.– Free ribosomes, on the other hand, make proteins

that stay local.• Key: Translation is slightly different for

ribosomes attached to the rough ER as compared to free ribosomes.

The Ultimate Goal• If a ribosome is producing a protein made for

secretion, the cell adds a signal peptide.– A signal peptide attaches to the forming polypeptide and

acts as…a signal.– Key: This occurs while the ribosome is making the

polypeptide – the process stops briefly.

The Ultimate Goal• A signal-recognition particle (SRP) can then interact

with the signal peptide and guide the ribosome to the ER.– This explains how the rough ER becomes rough.– Ribosomes move to it!

Practice

• Gene Expression – Translation POGIL– Up to second STOP sign – we’re going to go over

it!

The Big Picture

• “Label” everything you can by talking it out with your partner.

DNARNA

polymerase

RNA processing

Poly-A Tail

Amino acids

tRNA

Aminoacyl tRNA

synthetase

Large and small rRNA

subunits

E P Asites

Polypeptide

Nuclear pore5’ Cap

Semi-Closure

• We’re not done yet, but here’s a sort of philosophical thought for you:– “What is a gene?”

• Previously I’ve said it’s a stretch of DNA that codes for a particular protein, but that’s not entirely accurate.– Some genes code for multiple proteins.– Some genes code for RNA (not proteins).– Some genes overlap.– What about those promoters and control units and stuff?

The Currently-Accepted Definition

• According to your book:• “A gene is a region of DNA that can be

expressed to produce a final functional product that is either a polypeptide or an RNA molecule.”

Genes and Proteins

• Let’s review something else too – the idea that humans make around 100,000 proteins from 25,000 genes. How?– Different combinations of introns and exons.– The protein domains (the exons that are

combined) may be combined in different ways.– Post-translational modification can change the

protein in different ways.• Those are important concepts! ;)

Aside: The Longest Protein/Word

• The longest protein known to us right now is called Titin in short.• It’s got between 27,000 and 33,000 individual amino acids on it.• Multiply this by three to get a rough estimate of how many

mRNA nucleotides are involved.• What does it do?

– Helps you contract muscles.• And now…the full chemical name, which…again…was shortened

to “Titin.”– Note: Next slide in size 1.9 font.– http://www.sarahmcculloch.com/luminary-uprise/2009/longest-word/

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sparaginylalanylthreonylleucylarginylalanylphenylalanylalanylalanylglycylvalylthreonylprolylalanylgluta minylcysteinylphenylalanylglutamylmethionylleucylalanylleucylisoleucylarginylglutaminyllysylhistidylpr olylthreonylisoleucylprolylisoleucylglycylleucylleucylmethionyltyrosylalanylasparaginylleucylvalylphen ylalanylasparaginyllysylglycylisoleucylaspartylglutamylphenylalanyltyrosylalanylglutaminylcysteinylglu tamyllysylvalylglycylvalylaspartylserylvalylleucylvalylalanylaspartylvalylprolylvalylglutaminylglutamylse rylalanylprolylphenylalanylarginylglutaminylalanylalanylleucylarginylhistidylasparaginylvalylalanylproly lisoleucylphenylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylar ginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylargin ylalanylglycylvalylthreonylglycylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparagi nylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylgluta minylglycylphenylalanylglycylisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucyl aspartylalanylglycylalanylalanylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylis oleucylglutamylglutaminylhistidylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanyl alanylleucyllysylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetyls eryltyrosylserylisoleucylthreonylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylsery lvalyltryptophylalanylaspartylprolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylseryls erylleucylglycylasparaginylglutaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylt hreonylthreonylglutaminylvalylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprol ylphenylalanylprolylglutaminylserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysyl valyltyrosylarginyltyrosylasparaginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylle ucylglycylthreonylphenylalanylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalyl glutamylasparaginylglutaminylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspart ylalanylthreonylarginylarginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanyla sparaginylisoleucylasparaginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleuc yltyrosylasparaginylglutaminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucyl valyltryptophylthreonylserylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylp henylalanylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginyl glutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylala nylglycylvalylthreonylglycylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhi stidylleucylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminyl glycylphenylalanylglycylisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspa rtylalanylglycylalanylalanylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleuc ylglutamylglutaminylhistidylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanyl leucyllysylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyr osylserylisoleucylthreonylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalylt ryptophylalanylaspartylprolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylle ucylglycylasparaginylglutaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreo nylthreonylglutaminylvalylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphe nylalanylprolylglutaminylserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalylt yrosylarginyltyrosylasparaginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylg lycylthreonylphenylalanylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylgluta mylasparaginylglutaminylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylala nylthreonylarginylarginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylaspar aginylisoleucylasparaginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyr osylasparaginylglutaminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvaly ltryptophylthreonylserylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphen ylalanylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglut aminylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanyl glycylvalylthreonylglycylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhisti dylleucylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylgl ycylphenylalanylglycylisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspart ylalanylglycylalanylalanylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucyl glutamylglutaminylhistidylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylle ucyllysylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyro sylserylisoleucylthreonylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltry ptophylalanylaspartylprolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleu cylglycylasparaginylglutaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreony lthreonylglutaminylvalylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylpheny lalanylprolylglutaminylserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyro sylarginyltyrosylasparaginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglyc ylthreonylphenylalanylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamy lasparaginylglutaminylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylt hreonylarginylarginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparagin ylisoleucylasparaginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosyl asparaginylglutaminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryp tophylthreonylserylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylala nylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutamin ylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycyl valylthreonylglycylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleu cylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylph enylalanylglycylisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalany lglycylalanylalanylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutam ylglutaminylhistidylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllys ylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylseryl isoleucylthreonylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophyl alanylaspartylprolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycy lasparaginylglutaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreon ylglutaminylvalylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanyl prolylglutaminylserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylargi nyltyrosylasparaginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreo nylphenylalanylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylaspar aginylglutaminylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreon ylarginylarginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisol eucylasparaginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylaspar aginylglutaminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophy lthreonylserylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylal anylmethionylthreonylthreonylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylglutaminyll eucyllysylglutamylarginyllysylglutamylglycylalanylphenylalanylvalylprolylphenylalanylvalylthreonylleuc ylglycylaspartylprolylglycylisoleucylglutamylglutaminylserylleucyllysylisoleucylaspartylthreonylleucylis oleucylglutamylalanylglycylalanylaspartylalanylleucylglutamylleucylglycylisoleucylprolylphenylalanylse rylaspartylprolylleucylalanylaspartylglycylprolylthreonylisoleucylglutaminylasparaginylalanylthreonylle ucylarginylalanylphenylalanylalanylalanylglycylvalylthreonylprolylalanylglutaminylcysteinylphenylalany lglutamylmethionylleucylalanylleucylisoleucylarginylglutaminyllysylhistidylprolylthreonylisoleucylprolyli soleucylglycylleucylleucylmethionyltyrosylalanylasparaginylleucylvalylphenylalanylasparaginyllysylgly cylisoleucylaspartylglutamylphenylalanyltyrosylalanylglutaminylcysteinylglutamyllysylvalylglycylvalyla spartylserylvalylleucylvalylalanylaspartylvalylprolylvalylglutaminylglutamylserylalanylprolylphenylalany larginylglutaminylalanylalanylleucylarginylhistidylasparaginylvalylalanylprolylisoleucylphenylalanylisol eucylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylal anylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreony lglycylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalan yllysylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylgl ycylisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalany lalanylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminyl

histidylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphen ylalanylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylth reonylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspa rtylprolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparagin ylglutaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminy lvalylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylgluta minylserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosyl asparaginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenyl alanylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglut aminylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginyla rginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylaspa raginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglut aminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonyls erylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleu cylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalan ylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylgl ycylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyll ysylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglyc ylisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylal anylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhi stidylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenyl alanylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthre onylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspart ylprolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginyl glutaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylv alylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutami nylserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylas paraginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylal anylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylgluta minylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylar ginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylaspar aginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylgluta minylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylse rylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleuc ylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalany lseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylgly cylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanylly sylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycy lisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylala nylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhist idylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylal anylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreo nylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartyl prolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylgl utaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylval ylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminy lserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylaspar aginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalany laspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminy lglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylv alylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginyl leucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminyl asparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylala nylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcyst einylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylserylt yrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylala nylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleu cyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisole ucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylgl ycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidyla sparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylv alylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylser ylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolyli soleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutami nylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglut aminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylseryl threonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparagin ylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylasp artylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglut aminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalyl aspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleu cylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasp araginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanyl prolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylmethionylthreon ylthreonylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylglutaminylleucyllysylglutamylar ginyllysylglutamylglycylalanylphenylalanylvalylprolylphenylalanylvalylthreonylleucylglycylaspartylprolyl glycylisoleucylglutamylglutaminylserylleucyllysylisoleucylaspartylthreonylleucylisoleucylglutamylalanyl glycylalanylaspartylalanylleucylglutamylleucylglycylisoleucylprolylphenylalanylserylaspartylprolylleucy lalanylaspartylglycylprolylthreonylisoleucylglutaminylasparaginylalanylthreonylleucylarginylalanylphen ylalanylalanylalanylglycylvalylthreonylprolylalanylglutaminylcysteinylphenylalanylglutamylmethionylleu cylalanylleucylisoleucylarginylglutaminyllysylhistidylprolylthreonylisoleucylprolylisoleucylglycylleucylle ucylmethionyltyrosylalanylasparaginylleucylvalylphenylalanylasparaginyllysylglycylisoleucylaspartylgl utamylphenylalanyltyrosylalanylglutaminylcysteinylglutamyllysylvalylglycylvalylaspartylserylvalylleucyl valylalanylaspartylvalylprolylvalylglutaminylglutamylserylalanylprolylphenylalanylarginylglutaminylalan ylalanylleucylarginylhistidylasparaginylvalylalanylprolylisoleucylphenylalanylisoleucylcysteinylprolylpr olylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycyl arginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta myltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylala nylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylis oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla spartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalyla sparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylt hreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylala nyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylp rolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycy larginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta myltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylala nylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylis oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla spartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalyla sparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylt hreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylala nyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylp rolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycy larginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta myltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylala nylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylis oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla spartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalyla sparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylt hreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylala nyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylp rolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycy larginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta myltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylala nylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylis oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv

alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla spartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalyla sparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylt hreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylala nyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylmethionylthreonylthreon ylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylglutaminylleucyllysylglutamylarginyllys ylglutamylglycylalanylphenylalanylvalylprolylphenylalanylvalylthreonylleucylglycylaspartylprolylglycylis oleucylglutamylglutaminylserylleucyllysylisoleucylaspartylthreonylleucylisoleucylglutamylalanylglycylal anylaspartylalanylleucylglutamylleucylglycylisoleucylprolylphenylalanylserylaspartylprolylleucylalanyla spartylglycylprolylthreonylisoleucylglutaminylasparaginylalanylthreonylleucylarginylalanylphenylalanyl alanylalanylglycylvalylthreonylprolylalanylglutaminylcysteinylphenylalanylglutamylmethionylleucylalan ylleucylisoleucylarginylglutaminyllysylhistidylprolylthreonylisoleucylprolylisoleucylglycylleucylleucylmet hionyltyrosylalanylasparaginylleucylvalylphenylalanylasparaginyllysylglycylisoleucylaspartylglutamylp henylalanyltyrosylalanylglutaminylcysteinylglutamyllysylvalylglycylvalylaspartylserylvalylleucylvalylala nylaspartylvalylprolylvalylglutaminylglutamylserylalanylprolylphenylalanylarginylglutaminylalanylalanyl leucylarginylhistidylasparaginylvalylalanylprolylisoleucylphenylalanylisoleucylcysteinylprolylprolylaspa rtylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylgl ycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparagi nylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyros ylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolyl aspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylse rylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylgl utamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutaminylprolylm ethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylserylglutamin ylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylglutamylleuc ylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenylalanylgluta minylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglutaminylphe nylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylvalylarginylp henylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylvalylleucylasp artylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreonylarginylas paraginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylserylprolylthre onylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartylaspartylalany lthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalylasparaginylgl utamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylthreonylphe nylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylalanyltitinmethi onylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylprolylaspartyl alanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycy ltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparaginyl arginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosyla sparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolylasp artylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylserylg lycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylgluta 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glutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylserylglycy lserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylglutamyl prolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutaminylprolylmethion yllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylserylglutaminylphen ylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylglutamylleucylleuc ylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenylalanylglutaminylt hreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglutaminylphenylala nylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylvalylarginylphenyl alanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylvalylleucylaspartyl prolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreonylarginylaspara ginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylserylprolylthreonylt hreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartylaspartylalanylthre onylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalylasparaginylglutam ylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylthreonylphenylala nylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylalanyltitinmethionylgl utaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylprolylaspartylalanyl aspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyros ylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparaginylarginy lalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosylaspara ginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolylaspartylg lutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylserylglycyls erylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylglutamylpr olylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutaminylprolylmethionyll ysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylserylglutaminylphenyl alanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylglutamylleucylleucyl asparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenylalanylglutaminylthr eonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglutaminylphenylalan 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lylvalylglutaminylglutamylserylalanylprolylphenylalanylarginylglutaminylalanylalanylleucylarginylhistidy lasparaginylvalylalanylprolylisoleucylphenylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylas partylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonylt yrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparaginylarginylalanylalan ylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanyl alanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolylaspartylglutaminylv alyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylserylglycylserylalanyli soleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylglutamylprolylglutam yllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanyl alanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylserylglutaminylphenylalanylvaly lphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylglutamylleucylleucylasparagin ylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenylalanylglutaminylthreonylgluta minylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglutaminylphenylalanylserylgluta minylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylvalylarginylphenylalanylprolylgl ycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylvalylleucylaspartylprolylleucyliso leucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreonylarginylasparaginylarginyliso leucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylserylprolylthreonylthreonylalanylg lutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartylaspartylalanylthreonylvalylalanyli soleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalylasparaginylglutamylleucylvalylarg inylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylthreonylphenylalanylglutamylser ylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylalanyltitinmethionylglutaminylarginy ltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspart ylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonyltyros ylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparaginylarginylalanylalanylle ucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanylalan ylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolylaspartylglutaminylvalylly sylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylserylglycylserylalanylisole ucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylglutamylprolylglutamyllys ylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanylalan ylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylserylglutaminylphenylalanylvalylphe nylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylglutamylleucylleucylasparaginylval ylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenylalanylglutaminylthreonylglutamin ylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglutaminylphenylalanylserylglutamin ylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylvalylarginylphenylalanylprolylglycy laspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylvalylleucylaspartylprolylleucylisoleu cylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreonylarginylasparaginylarginylisoleu cylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylserylprolylthreonylthreonylalanylglut amylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartylaspartylalanylthreonylvalylalanyliso leucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalylasparaginylglutamylleucylvalylargin ylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylthreonylphenylalanylglutamylseryl methionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylalanyltitinmethionylglutaminylarginylt yrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspartyl aspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosyl leucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparaginylarginylalanylalanylleuc ylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanylalanyl 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lglycylisoleucylprolylphenylalanylserylaspartylprolylleucylalanylaspartylglycylprolylthreonylisoleucylglu taminylasparaginylalanylthreonylleucylarginylalanylphenylalanylalanylalanylglycylvalylthreonylprolylal anylglutaminylcysteinylphenylalanylglutamylmethionylleucylalanylleucylisoleucylarginylglutaminyllysyl histidylprolylthreonylisoleucylprolylisoleucylglycylleucylleucylmethionyltyrosylalanylasparaginylleucylv alylphenylalanylasparaginyllysylglycylisoleucylaspartylglutamylphenylalanyltyrosylalanylglutaminylcys teinylglutamyllysylvalylglycylvalylaspartylserylvalylleucylvalylalanylaspartylvalylprolylvalylglutaminylgl utamylserylalanylprolylphenylalanylarginylglutaminylalanylalanylleucylarginylhistidylasparaginylvalylal anylprolylisoleucylphenylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleuc ylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucyl 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ylserylleucylphenylalanylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleuc ylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucyl serylarginylalanylglycylvalylthreonylglycylalanylglutamylasparaginylarginylalanylalanylleucylprolylleuc ylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolyll eucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalan ylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysyli soleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylglutamylprolylglutamyllysylmethionylle ucylalanylalanylleucyllysylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanylalanylthreonylargi nylacetylseryltyrosylserylisoleucylthreonylserylprolylserylglutaminylphenylalanylvalylphenylalanylleuc 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ylaspartylthreonylleucylisoleucylglutamylalanylglycylalanylaspartylalanylleucylglutamylleucylglycylisol eucylprolylphenylalanylserylaspartylprolylleucylalanylaspartylglycylprolylthreonylisoleucylglutaminylas paraginylalanylthreonylleucylarginylalanylphenylalanylalanylalanylglycylvalylthreonylprolylalanylgluta minylcysteinylphenylalanylglutamylmethionylleucylalanylleucylisoleucylarginylglutaminyllysylhistidylpr olylthreonylisoleucylprolylisoleucylglycylleucylleucylmethionyltyrosylalanylasparaginylleucylvalylphen ylalanylasparaginyllysylglycylisoleucylaspartylglutamylphenylalanyltyrosylalanylglutaminylcysteinylglu tamyllysylvalylglycylvalylaspartylserylvalylleucylvalylalanylaspartylvalylprolylvalylglutaminylglutamylse rylalanylprolylphenylalanylarginylglutaminylalanylalanylleucylarginylhistidylasparaginylvalylalanylproly lisoleucylphenylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylar 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erylleucylglycylasparaginylglutaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylt hreonylthreonylglutaminylvalylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprol ylphenylalanylprolylglutaminylserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysyl valyltyrosylarginyltyrosylasparaginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylle ucylglycylthreonylphenylalanylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalyl glutamylasparaginylglutaminylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspart ylalanylthreonylarginylarginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanyla sparaginylisoleucylasparaginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleuc yltyrosylasparaginylglutaminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucyl 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asparaginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenyl alanylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglut aminylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginyla rginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylaspa raginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglut aminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonyls erylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleu cylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalan ylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylgl 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ylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalany lseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylgly cylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanylly sylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycy lisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylala nylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhist idylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylal anylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreo nylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartyl prolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylgl utaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylval ylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminy lserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylaspar aginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalany laspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminy lglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylv alylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginyl leucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminyl asparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylala nylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcyst einylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylserylt yrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylala nylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleu cyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisole ucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylgl ycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidyla sparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylv alylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylser ylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolyli soleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutami nylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglut aminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylseryl threonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparagin ylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylasp artylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglut aminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalyl aspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleu cylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasp araginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanyl prolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylmethionylthreon ylthreonylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylglutaminylleucyllysylglutamylar ginyllysylglutamylglycylalanylphenylalanylvalylprolylphenylalanylvalylthreonylleucylglycylaspartylprolyl glycylisoleucylglutamylglutaminylserylleucyllysylisoleucylaspartylthreonylleucylisoleucylglutamylalanyl glycylalanylaspartylalanylleucylglutamylleucylglycylisoleucylprolylphenylalanylserylaspartylprolylleucy lalanylaspartylglycylprolylthreonylisoleucylglutaminylasparaginylalanylthreonylleucylarginylalanylphen 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valylalanylaspartylvalylprolylvalylglutaminylglutamylserylalanylprolylphenylalanylarginylglutaminylalan ylalanylleucylarginylhistidylasparaginylvalylalanylprolylisoleucylphenylalanylisoleucylcysteinylprolylpr olylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycyl arginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta myltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylala nylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylis oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla 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oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla spartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalyla sparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylt hreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylala nyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylp rolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycy larginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta 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larginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta myltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylala nylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylis oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla spartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalyla sparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylt hreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylala nyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylmethionylthreonylthreon ylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylglutaminylleucyllysylglutamylarginyllys ylglutamylglycylalanylphenylalanylvalylprolylphenylalanylvalylthreonylleucylglycylaspartylprolylglycylis oleucylglutamylglutaminylserylleucyllysylisoleucylaspartylthreonylleucylisoleucylglutamylalanylglycylal anylaspartylalanylleucylglutamylleucylglycylisoleucylprolylphenylalanylserylaspartylprolylleucylalanyla spartylglycylprolylthreonylisoleucylglutaminylasparaginylalanylthreonylleucylarginylalanylphenylalanyl alanylalanylglycylvalylthreonylprolylalanylglutaminylcysteinylphenylalanylglutamylmethionylleucylalan ylleucylisoleucylarginylglutaminyllysylhistidylprolylthreonylisoleucylprolylisoleucylglycylleucylleucylmet hionyltyrosylalanylasparaginylleucylvalylphenylalanylasparaginyllysylglycylisoleucylaspartylglutamylp henylalanyltyrosylalanylglutaminylcysteinylglutamyllysylvalylglycylvalylaspartylserylvalylleucylvalylala nylaspartylvalylprolylvalylglutaminylglutamylserylalanylprolylphenylalanylarginylglutaminylalanylalanyl leucylarginylhistidylasparaginylvalylalanylprolylisoleucylphenylalanylisoleucylcysteinylprolylprolylaspa rtylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylgl ycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparagi nylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyros ylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolyl aspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylse rylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylgl utamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutaminylprolylm ethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylserylglutamin ylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylglutamylleuc ylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenylalanylgluta minylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglutaminylphe nylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylvalylarginylp henylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylvalylleucylasp artylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreonylarginylas paraginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylserylprolylthre onylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartylaspartylalany lthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalylasparaginylgl utamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylthreonylphe nylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylalanyltitinmethi onylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylprolylaspartyl alanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycy ltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparaginyl arginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosyla sparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolylasp 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aspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyros ylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparaginylarginy lalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosylaspara ginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolylaspartylg

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partylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonylt yrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparaginylarginylalanylalan ylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanyl alanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolylaspartylglutaminylv alyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylserylglycylserylalanyli soleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylglutamylprolylglutam yllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanyl alanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylserylglutaminylphenylalanylvaly lphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylglutamylleucylleucylasparagin 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ylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylalanyltitinmethionylglutaminylarginy ltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspart ylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonyltyros ylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamylasparaginylarginylalanylalanylle ucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanylalan ylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolylaspartylglutaminylvalylly sylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylserylglycylserylalanylisole ucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylglutamylprolylglutamyllys ylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanylalan 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ylisoleucylasparaginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosyl asparaginylglutaminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryp tophylthreonylserylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylala nylalanylisoleucylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutamin ylisoleucylalanylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycyl valylthreonylglycylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleu cylvalylalanyllysylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylph enylalanylglycylisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalany lglycylalanylalanylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutam ylglutaminylhistidylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllys ylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylseryl isoleucylthreonylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophyl alanylaspartylprolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycy lasparaginylglutaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreon ylglutaminylvalylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanyl prolylglutaminylserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylargi nyltyrosylasparaginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreo nylphenylalanylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylaspar aginylglutaminylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreon ylarginylarginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisol eucylasparaginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylaspar aginylglutaminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophy lthreonylserylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylal anylmethionylthreonylthreonylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylglutaminyll eucyllysylglutamylarginyllysylglutamylglycylalanylphenylalanylvalylprolylphenylalanylvalylthreonylleuc ylglycylaspartylprolylglycylisoleucylglutamylglutaminylserylleucyllysylisoleucylaspartylthreonylleucylis oleucylglutamylalanylglycylalanylaspartylalanylleucylglutamylleucylglycylisoleucylprolylphenylalanylse rylaspartylprolylleucylalanylaspartylglycylprolylthreonylisoleucylglutaminylasparaginylalanylthreonylle ucylarginylalanylphenylalanylalanylalanylglycylvalylthreonylprolylalanylglutaminylcysteinylphenylalany lglutamylmethionylleucylalanylleucylisoleucylarginylglutaminyllysylhistidylprolylthreonylisoleucylprolyli soleucylglycylleucylleucylmethionyltyrosylalanylasparaginylleucylvalylphenylalanylasparaginyllysylgly cylisoleucylaspartylglutamylphenylalanyltyrosylalanylglutaminylcysteinylglutamyllysylvalylglycylvalyla spartylserylvalylleucylvalylalanylaspartylvalylprolylvalylglutaminylglutamylserylalanylprolylphenylalany larginylglutaminylalanylalanylleucylarginylhistidylasparaginylvalylalanylprolylisoleucylphenylalanylisol eucylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylal anylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreony lglycylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalan yllysylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylgl ycylisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalany lalanylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminyl histidylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphen ylalanylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylth reonylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspa rtylprolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparagin ylglutaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminy lvalylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylgluta minylserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosyl asparaginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenyl alanylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglut

aminylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginyla rginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylaspa raginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglut aminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonyls erylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleu cylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalan ylseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylgl ycylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyll ysylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglyc ylisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylal anylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhi stidylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenyl alanylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthre onylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspart ylprolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginyl glutaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylv alylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutami nylserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylas paraginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylal anylaspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylgluta minylglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylar ginylvalylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylaspar aginylleucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylgluta minylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylse rylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleuc ylcysteinylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalany lseryltyrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylgly cylalanylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanylly sylleucyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycy lisoleucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylala nylglycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhist idylasparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylal anylvalylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreo nylserylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartyl prolylisoleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylgl utaminylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylval ylglutaminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminy lserylthreonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylaspar aginylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalany laspartylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminy lglutaminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylv alylaspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginyl leucylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminyl asparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylala nylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcyst einylprolylprolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylserylt yrosylglycylarginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylala nylglutamylasparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleu cyllysylglutamyltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisole ucylserylalanylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylgl ycylalanylisoleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidyla sparaginylisoleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylv alylglutaminylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylser ylprolylserylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolyli soleucylglutamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutami nylphenylalanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglut aminylglutaminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylseryl threonylvalylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparagin ylalanylvalylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylasp artylthreonylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglut aminylserylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalyl aspartylaspartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleu cylvalylasparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasp araginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanyl prolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylmethionylthreon ylthreonylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylglutaminylleucyllysylglutamylar ginyllysylglutamylglycylalanylphenylalanylvalylprolylphenylalanylvalylthreonylleucylglycylaspartylprolyl glycylisoleucylglutamylglutaminylserylleucyllysylisoleucylaspartylthreonylleucylisoleucylglutamylalanyl glycylalanylaspartylalanylleucylglutamylleucylglycylisoleucylprolylphenylalanylserylaspartylprolylleucy lalanylaspartylglycylprolylthreonylisoleucylglutaminylasparaginylalanylthreonylleucylarginylalanylphen ylalanylalanylalanylglycylvalylthreonylprolylalanylglutaminylcysteinylphenylalanylglutamylmethionylleu cylalanylleucylisoleucylarginylglutaminyllysylhistidylprolylthreonylisoleucylprolylisoleucylglycylleucylle ucylmethionyltyrosylalanylasparaginylleucylvalylphenylalanylasparaginyllysylglycylisoleucylaspartylgl utamylphenylalanyltyrosylalanylglutaminylcysteinylglutamyllysylvalylglycylvalylaspartylserylvalylleucyl valylalanylaspartylvalylprolylvalylglutaminylglutamylserylalanylprolylphenylalanylarginylglutaminylalan ylalanylleucylarginylhistidylasparaginylvalylalanylprolylisoleucylphenylalanylisoleucylcysteinylprolylpr olylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycyl arginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta myltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylala nylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylis oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla spartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalyla sparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylt hreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylala nyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylp rolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycy larginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta myltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylala nylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylis oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla spartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalyla sparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylt hreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylala nyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylp rolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycy larginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta myltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylala nylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylis oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla spartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalyla sparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylt hreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylala nyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylisoleucylcysteinylprolylp rolylaspartylalanylaspartylaspartylaspartylleucylleucylarginylglutaminylisoleucylalanylseryltyrosylglycy larginylglycyltyrosylthreonyltyrosylleucylleucylserylarginylalanylglycylvalylthreonylglycylalanylglutamyl asparaginylarginylalanylalanylleucylprolylleucylasparaginylhistidylleucylvalylalanyllysylleucyllysylgluta myltyrosylasparaginylalanylalanylprolylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylala nylprolylaspartylglutaminylvalyllysylalanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylis oleucylserylglycylserylalanylisoleucylvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginyli soleucylglutamylprolylglutamyllysylmethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutami nylprolylmethionyllysylalanylalanylthreonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylser ylglutaminylphenylalanylvalylphenylalanylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylgl utamylleucylleucylasparaginylvalylcysteinylthreonylserylserylleucylglycylasparaginylglutaminylphenyl alanylglutaminylthreonylglutaminylglutaminylalanylarginylthreonylthreonylglutaminylvalylglutaminylglu taminylphenylalanylserylglutaminylvalyltryptophyllysylprolylphenylalanylprolylglutaminylserylthreonylv alylarginylphenylalanylprolylglycylaspartylvalyltyrosyllysylvalyltyrosylarginyltyrosylasparaginylalanylva lylleucylaspartylprolylleucylisoleucylthreonylalanylleucylleucylglycylthreonylphenylalanylaspartylthreo nylarginylasparaginylarginylisoleucylisoleucylglutamylvalylglutamylasparaginylglutaminylglutaminylse rylprolylthreonylthreonylalanylglutamylthreonylleucylaspartylalanylthreonylarginylarginylvalylaspartyla spartylalanylthreonylvalylalanylisoleucylarginylserylalanylasparaginylisoleucylasparaginylleucylvalyla sparaginylglutamylleucylvalylarginylglycylthreonylglycylleucyltyrosylasparaginylglutaminylasparaginylt hreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreonylserylalanylprolylala nyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylmethionylthreonylthreon ylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylglutaminylleucyllysylglutamylarginyllys ylglutamylglycylalanylphenylalanylvalylprolylphenylalanylvalylthreonylleucylglycylaspartylprolylglycylis oleucylglutamylglutaminylserylleucyllysylisoleucylaspartylthreonylleucylisoleucylglutamylalanylglycylal anylaspartylalanylleucylglutamylleucylglycylisoleucylprolylphenylalanylserylaspartylprolylleucylalanyla spartylglycylprolylthreonylisoleucylglutaminylasparaginylalanylthreonylleucylarginylalanylphenylalanyl alanylalanylglycylvalylthreonylprolylalanylglutaminylcysteinylphenylalanylglutamylmethionylleucylalan 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glutaminylasparaginylthreonylphenylalanylglutamylserylmethionylserylglycylleucylvalyltryptophylthreo nylserylalanylprolylalanyltitinmethionylglutaminylarginyltyrosylglutamylserylleucylphenylalanylalanylpr olylprolylleucylglutaminylglycylphenylalanylglycylisoleucylserylalanylprolylaspartylglutaminylvalyllysyl alanylalanylisoleucylaspartylalanylglycylalanylalanylglycylalanylisoleucylserylglycylserylalanylisoleucy lvalyllysylisoleucylisoleucylglutamylglutaminylhistidylasparaginylisoleucylglutamylprolylglutamyllysylm ethionylleucylalanylalanylleucyllysylvalylphenylalanylvalylglutaminylprolylmethionyllysylalanylalanylthr eonylarginylacetylseryltyrosylserylisoleucylthreonylserylprolylserylglutaminylphenylalanylvalylphenylal anylleucylserylserylvalyltryptophylalanylaspartylprolylisoleucylglutamylleucylleucylasparaginylvalylcys teinylthreonylserylserylleucylglycylasparaginylglutaminylphenylalanylglutaminylthreonylglutaminylglut 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Wow.

• We’ve covered a lot.• We’ve learned how to go from DNA to mRNA and

how to go from mRNA to protein.• The only thing left in the Central Dogma is how

to go from proteins to traits, so let’s do that now.• Oh wait…we already did. Remember Mendelian

genetics?– So in a weird sense, only just now have we covered

the basics of biology.

A Broad Review: Prokaryotes• Gene Regulation

– Operons• Promoter• Operator• Repressor• Genes

• Transcription– Initiation: RNA polymerase attaches to DNA.– Elongation: RNA polymerase makes mRNA by reading DNA.– Termination: Termination sequence releases RNA polymerase and mRNA.

• Translation– Initiation: Ribosomal subunits assemble around mRNA.– Elongation: Ribosome makes polypeptides using tRNA [A, P, E sites]. – Termination: Stop codon imports release factor, ribosome disassembles.

A Broad Review: Eukaryotes• Gene Regulation

– Activators.– Promotors.– Transcription factors.

• Transcription– Initiation: Transcription factors attach to TATA box. RNA polymerase II attaches to

transcription factors.– Elongation: RNA polymerase II makes mRNA by reading DNA.– Termination: Termination sequence immediately releases mRNA and eventually

releases RNA polymerase II.• RNA Processing

– Add 5’ cap and poly-A tail.– Remove introns.

• Translation– Initiation: Ribosomal subunits assemble around mRNA.– Elongation: Ribosome makes polypeptides using tRNA [A, P, E sites]. – Termination: Stop codon imports release factor, ribosome disassembles.

So uh…

• …what’s left?• Oh, I know.• Evolution.– …er…rather…mutation.

• What happens when something goes wrong?

Mutations

• We’ve already gone over chromosomal mutations (the really large scale ones like nondisjunction, deletion, et cetera).

• Now we look at a much smaller scale mutation, one that is within the genetic code.

• We call these point mutations as they act on a single nucleotide location.– A point mutation, or just a difference of a letter in

general, is sometimes called a SNP (single nucleotide polymorphism).

Point Mutations

• There are generally four effects to point mutations:– A missense mutation makes a different amino acid.– A nonsense mutation makes a stop codon when there

shouldn’t be one.– A neutral or silent mutation has no effect.• Remember that whole codon redundancy/wobble thing?

– A frame-shift mutation is a large-scale mutation that could cause any combination of the above.

Missense Mutation Example

• The mutation that causes sickle-cell anemia is a missense mutation that change glutamate to valine:

Frame-Shift Mutations

• Here’s a sample sentence:– THE RED DOG ATE THE BIG BUG

• The spaces provide a reading frame.• Here’s a sample mRNA strand:– AUG CCC CUU UGA

• The spaces provide a reading frame.• But what if you accidentally delete a letter?– THR EDD OGA TET HEB IBU G– AUC CCC UUU GA

Frame-Shift Mutations

• As you might guess, a frame-shift mutation is generally bad news.

• All codons downstream of the mutation are affected.

• Let’s test our ability to spot frame-shift mutations as well as point mutations in the following sample RNA strands.– Grab your universal genetic codes…

AUG UUU GAG UGA UCC

• Methionine• Phenylalanine• Glutamate• [Stop]• [Serine]

AUG UUU GAG UGA UCC

• Mutation 1: What if I add a C here?– This is an insertion mutation.

• AUG UUU CGA GUG AUC C• What happens?– Methionine– Phenylalanine– Arginine– Valine– Isoleucine

• No stop codon!• This is a frame-shift mutation and a missense mutation.

AUG UUU GAG UGA UCC

• Mutation 2: What if I add a GA here?– This is a frame-shift mutation.

• AUG UGA UUG AGU GAU CC• What happens?– Methionine– [Stop]

• The early stop codon prevents the full protein from forming.

• This is a nonsense mutation.

AUG UUU GAG UGA UCC

• Mutation 3: What if I delete this A?– This is a deletion mutation.

• AUG UUU GAG UGU CC• What happens?– Methionine– Phenylalanine– Glutamate– Cysteine

• No stop codon expressed - protein keeps going!

AUG UUU GAG UGA UCC

• Mutation 4: What if I change this A to a G?– This is a substitution mutation.

• AUG UUU GAG UGG UCC– Methionine– Phenylalanine– Glutamate– Tryptophan– Serine

• No stop codon expressed - protein keeps going!

AUG UUU GAG UGA UCC

• Mutation 5: What if I change this U to a C?– This is a substitution mutation.

• ACG UUU GAG UGA UCC• No start codon expressed - protein never formed!

AUG UUU GAG UGA UCC

• Mutation 6: What if I change this A to a U?– This is a substitution mutation.

• AUG UUU GUG UGA UCC– Methionine– Phenylalanine– Valine– [Stop]– [Serine]

• Glutamate is changed to valine.• Not a frame-shift mutation (is a missense).– Unfortunately, this is all it takes to get sickle-cell anemia.

Sickle-Cell Anemia

• That’s the effect a single nucleotide change can have…

http://adultstemcellawareness.files.wordpress.com/2007/10/sickle-cell.jpg

AUG UUU GAG UGA UCC

• Mutation 7: What if I change this U to a C?– This is a substitution mutation.

• AUG UUC GAG UGA UCC– Methionine– Phenylalanine– Glutamate– [Stop]– [Serine]

• No change - this is a neutral mutation!

Closure

• So here’s a question.• Sickle-cell anemia, as we just said, is a disorder

caused by a single-base mutation in hemoglobin genes that changes a glutamate amino acid to valine.

• For individuals with sickle-cell anemia, is the mutation in their DNA or in their RNA?– DNA. It’s a lifelong mutation; RNA is temporary.

Closure Explanation

• It’s like this:– You make a cake and the recipe calls for 2 sticks of

butter.– You put in 20. What happens?

• Your cake tastes like poo.

– When you redo it, you’ll probably be fine.– But, if the recipe has a typo in it, no matter how

many times you make the cake, it’ll taste like poo.• DNA mutations are like having typos in the

recipe; RNA mutations are like screwing up in an “isolated incident.”

Closure Deux

• Gene Expression – Translation POGIL– Finish it – we’re going to go over it!