Chapter 24: Nucleic Acid Structure Flashcards

1
Q

ultimate source of biological information?

A

Nucleic acids

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2
Q

The shapes and activities of individual cells are, to a large extent, determined by genetic instructions contained in

A

DNA (or RNA, in some viruses).

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3
Q

According to the central dogma of molecular biology, sequences of nucleotide bases in DNA encode?

A

the amino acid sequences of proteins.

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4
Q

What are the two kinds of nucleic acids and what do they do?

A

store information and make it available to the cell.

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5
Q

The structures of DNA and RNA are consistent with what 4 things?

A
  1. Genetic information must be stored in a form that is manageable in size and stable over a long period.
  2. Genetic information must be decoded—often many times—in order to be used. Transcription is the process by which nucleotide sequences in DNA are copied onto RNA so that they can direct protein synthesis, a process known as translation.
  3. Information contained in DNA or RNA must be accessible to proteins and other nucleic acids. These agents must recognize nucleic acids (in many cases, in a sequence-specific fashion) and bind to them in a way that alters their function.
  4. The progeny of an organism must be equipped with the same set of instructions as in the parent. Thus, DNA is replicated (an exact copy made) so that each daughter cell receives the same information.
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6
Q

What is the structural difference between thymine and uracil?

A

The only difference between them is the presence of an extra methyl group in the thymine structure.

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7
Q

Which form of DNA is biologically most common?

A

B-DNA

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8
Q

Structural features of B-DNA?

A
  1. The two antiparallel polynucleotide strands wind in a right-handed manner around a common axis to produce an ∼20-Å-diameter double helix.
  2. The planes of the nucleotide bases, which form hydrogen-bonded pairs, are nearly perpendicular to the helix axis. In B-DNA, the bases occupy the core of the helix while the sugar–phosphate backbones wind around the outside, forming the major and minor grooves. Only the edges of the base pairs are exposed to solvent.
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9
Q

What accounts for the near perfect symmetry of DNA?

A

Each base pair has approximately the same width (Fig. 24-1), which accounts for the near-perfect symmetry of the DNA molecule, regardless of base composition. A ∙ T and G ∙ C base pairs are interchangeable: They can replace each other in the double helix without altering the positions of the sugar–phosphate backbones’ C1′ atoms. Likewise, the partners of a Watson–Crick base pair can be switched (i.e., by changing a G ∙ C to a C ∙ G or an A ∙ T to a T ∙ A). In contrast, any other combination of bases would significantly distort the double helix.

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10
Q

how many bp per turn of B DNA and helical twist?

A

The canonical (ideal) B-DNA helix has 10 base pairs (bp) per turn (a helical twist of 36° per bp) and, since the aromatic bases have van der Waals thicknesses of 3.4 Å and are partially stacked on each other, the helix has a pitch (rise per turn) of 34 Å.

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11
Q

The major structural variants of B-DNA?

A

A-DNA and Z-DNA

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12
Q

Describe A-DNA

A

A-DNA’s Base Pairs Are Inclined to the Helix Axis.

Under dehydrating conditions, B-DNA undergoes a reversible conformational change to A-DNA, which forms a wider and flatter right-handed helix than does B-DNA.

Has an an axial hole

A-DNA’s most striking feature, however, is that the planes of its base pairs are tilted 20° with respect to the helix axis. Since the axis does not pass through its base pairs, A-DNA has a deep major groove and a very shallow minor groove; it can be described as a flat ribbon wound around a 6-Å-diameter cylindrical hole.

11bp

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13
Q

Where does A-DNA occur?

A

A short segment of A-DNA occurs in the active site of DNA polymerase during the replication of DNA

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14
Q

Describe Z-DNA

A

Z-DNA Forms a Left-Handed Helix.

a deep minor groove, and no discernible major groove.

formed under high salt concentration. Left-handed, 12 base pairs per turn, a pitch of 45 Å.

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15
Q

Describe Z-DNA

A

Z-DNA Forms a Left-Handed Helix.

a deep minor groove, and no discernible major groove.

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16
Q

Does Z-DNA have a biological function?

A

-The discovery of Z-DNA–binding proteins strongly suggests that Z-DNA does exist in vivo.

-One Zα domain binds to each strand of Z-DNA via hydrogen bonds and ionic interactions between polar and basic side chains and the sugarphosphate backbone of the DNA; none of the DNA’s bases participate in these associations

-Sequences capable of forming Z-DNA frequently occur near the start of genes, and the reversible conversion of B-DNA to Z-DNA at these sites may play a role in the control of transcription.

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17
Q

Are A:T and G:C base pairs are interchangeable? Why?

A

Yes, They can replace each other in the double helix without altering the positions of the sugar–phosphate backbones’ C1′ atoms.

This is because each base pair has approximately the same width

In contrast, any other combination of bases would significantly distort the double helix.

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18
Q

The major structural features of three types of DNA

A

A-DNA and B-DNA are right handed helixes & anti conformation
Z-DNA is left handed helix. Anti for pyrimidines; syn for purines

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19
Q

do the structurally distinct A, B, and Z forms of DNA freely interconvert in vivo?

A

No, Rather, the transition from one form to another requires unusual physical conditions (e.g., dehydration) or the influence of DNA-binding proteins.

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20
Q

Real DNA molecules deviate from the ideal structures, expand on this. Why is this important for proteins.

A

X-Ray structures of B-DNA segments reveal that individual residues significantly depart from the average conformation in a sequence-dependent manner.
For example, the helical twist per base pair may range from 26° to 43°.

Each base pair can also deviate from its ideal conformation by rolling or twisting like the blade of a propeller.

Such conformational variation appears to be important for the sequence-specific recognition of DNA by the proteins that process genetic information.

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21
Q

DNA has structural flexibility but the flexibility is limited. Why is DNA flexible?

A

it is imperative that the molecules be somewhat flexible so that they can be packaged in cells. DNA helices can adopt different degrees of curvature ranging from gentle arcs to sharp bends. The more severe distortions from linearity generally occur in response to the binding of specific proteins.

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22
Q

Conformation of a nucleotide is specified by ?

A

The six torsion angles of the sugar-phosphate backbone and the torsion angle of the glycosidic bond connecting base and sugar.

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23
Q

Are the six torsion angles of the sugar-phosphate backbone and the torsion angle around the glycosidic bond restricted? Due to what?

A

Yes, due to internal constraints that restrict their rotational freedom.

Because of non-covalent interactions between the ribose ring and the phosphate groups, and steric interference between residues.

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24
Q

The two most common ribose conformations are known as ?

A

C3′-endo and C2′-endo

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25
Q

The flexibility of the ribose ring is also limited.
The two most common ribose conformations are known as C3′-endo and C2′-endo; “endo” (Greek: endon, within) indicates that the displaced atom is on the same side of the ring as C5′, whereas “exo” (Greek: exo, out of) indicates displacement toward the opposite side of the ring from C5′.

What are the endo conformations of the DNAs

A

The flexibility of the ribose ring is also limited.

  • B-DNA has the C2’-endo conformation.
  • A-DNA has the C3’-endo conformation.
  • Z-DNA has C3’-endo of purine nucleotides and C2’-endo of pyrimidine nucleotides.
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26
Q

The rotation of a base around its glycosidic bond (angle χ ) is greatly hindered. Purine residues have two sterically permissible orientations known as the

A

syn (Greek: with) and anti (Greek: against) conformations

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27
Q

Only the anti conformation of ____ is stable, because?

A

pyrimidines because in the syn conformation, the sugar residue sterically interferes with the pyrimidine’s C2 substituent.

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28
Q

In most double-helical nucleic acids, all bases are in the _____ conformation, which the exception of?

A

anti

The exception is Z-DNA (Section 24-1A), in which the alternating pyrimidine and purine residues are anti and syn, respectively (this is one reason why the repeating unit of Z-DNA is a dinucleotide).

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29
Q

The chromosomes of many viruses and bacteria are circular molecules of duplex DNA. These molecules are a result of?

A

DNA supercoiling.

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30
Q

Describe Supercoiled DNA
Why is it important

A

Supercoiled DNA molecules are more compact than “relaxed” molecules with the same number of nucleotides.

Supercoiling is important for packaging DNA in cells, and for unwinding DNA during replication and transcription.

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31
Q

The mathematically expressed topology of supercoiled DNA

A

L=T+W

L, the linking number, is the number of times that one DNA strand winds around the other.

T, the twist, is the number of complete revolutions that one polynucleotide strand makes around the duplex axis
T is positive for right-handed duplex turns so that, for B-DNA,

  1. W, the writhing number, is the number of turns that the
    W, the writhing number, is the number of turns that the duplex axis makes around the superhelix axis. It is a measure of the DNA’s superhelicity.
32
Q

How to calculate the twist number in B-DNA? What If circular DNA helix lies flat on a plane?

A

the twist is normally the number of base pairs divided by 10.5 (the observed number of base pairs per turn of the B-DNA double helix in aqueous solution).
If circular DNA helix lies flat on a plane, the twist number is equal to the linking number.

33
Q

Supercoiled DNA Is Relaxed by?

A

Nicking One Strand by an enzyme pancreatic dnase I that cleaves phosphodiester bonds. , which cleaves only one strand of a duplex DNA..

  • Topoisomerases control DNA supercoiling by making breaks in DNA double helix.
34
Q

What are Topoisomerases? When is it used?

A

control the level of supercoiling in DNA, are so named because they alter DNA’s topological state (linking number) but not its covalent structure. for replication and trancription unwinding

35
Q

Two classes of topoisomerases in prokaryotes and eukaryotes?

A

Type I topoisomerases create transient single-strand breaks in DNA, including Type 1A and Type 1B;

Type II topoisomerases make transient double-strand breaks in DNA.

36
Q

Why must naturally occurring DNAs supercoiling be negative.

A

In such basic biological processes as replication and transcription, complementary polynucleotide strands must separate. The negative supercoiling of naturally occurring DNAs in both prokaryotes and eukaryotes promotes such separations since it tends to unwind the duplex helix

37
Q

Type I topoisomerase function? How do they relax and unwind DNA?

Which amino acid in the active site of topoisomerases acts to cleave DNA starnds and how?

A

Type IA topoisomerases catalyze the relaxation of negative supercoils in DNA by increasing its linking number in increments of one turn

Type IB only occur in eukaryotic, which can relax both negative and positive supercoils in DNA.
Type IB Topoisomerases Relax Supercoiled DNA via Controlled Rotation.

  • Type I topoisomerases cut single-strand DNA, pass a single-stranded loop through the resulting gap, and then reseal the break.
  • Type IA unwinds supercoiled DNA via the strand passage mechanism. The active site Tyr attacks the phosphate group forming a Tyr-5’-P and a free 3’-OH of the cleaved strand
  • Type IB active site Tyr attacks the phosphate group forming a covalent linkage with the 3” end of the cleaved strand and a free 5’-OH. Type IB relaxes supercoiled DNA using a controlled rotation mechanism.
38
Q

Type II topoisomerase function?

A

Type II are multimeric enzymes that require ATP hydrolysis to complete a reaction cycle in which two DNA strands are cleaved.

  • Both prokaryotic and eukaryotic topo II relax negative and positive supercoils in DNA, but only prokaryotic topo II (also known as DNA gyrase) can introduce negative supercoils.
  • Topo II has a pair of catalytic Tyr residues in its active site, each mediates a break in one strand of DNA double helix, creating a double-strand breaks.
  • Topo II mediate cleavage of the two DNA strands at staggered sites to produce four-base sticky ends.
  • Gyrases are A2B2 heterotetramers, B subunit hydrolyzes ATP and A subunit is the catalytic subunit, while eukaryotic Topo II has fused A and B to form a homodimer

requires the input of free energy

39
Q

Explain why most nucleotides adopt the anti conformation.

A

interference with pyrimidines c2

40
Q

A double-stranded nucleic acid structure is stabilized by what 3 events? Which is the most stable?

A

base pairing, stacking interactions, and by ionic interactions.

Watson–Crick geometry
is the most stable mode of base pairing in the double helix, even though non-WatsonCrick base pairs are theoretically possible.

A:T, T:A, G:C, and C:G base pairs have same width and is geometrically similar, but other base pairs do not.

41
Q

Why do non-Watson Crick base pairs not occur as much?

A

the bases in a Watson–Crick pair have a higher mutual affinity than those in a non-Watson–Crick pair.

42
Q

Which event contributes little to the stability of nucleic acid structure.

A

It is clear that hydrogen bonding is required for the specificity of base pairing in DNA. Yet, as is also true for proteins (Section 6-4A), hydrogen bonding contributes little to the stability of nucleic acid structures. they get easily replaced during denaturation

43
Q

What are the Base stacking interactions?
What do they result from? Which base pair has greater stacking interactions?

A

the layering of bases on top of each other in the double-stranded structure. Base stacking is thermodynamically favorable because of the hydrophobic effect and because of van der Waals interactions between stacked bases

Stacking Interactions Result from Hydrophobic Forces. Purines and pyrimidines tend to form extended stacks of planar parallel molecules.

These stacking interactions are a form of van der Waals interaction.

Interactions between stacked G and C bases are greater than those between stacked A and T bases, which largely accounts for the greater thermal stability and Tm of DNAs with a high G + C content.

44
Q

The stacking interactions between base pairs are a form of what?

A

These stacking interactions are a form of van der Waals interaction

45
Q

Describe the Ionic interactions that stabilize nucleic acids.

A

Cations can shield the negative charge of the phosphate groups in DNA molecule and affect the stability of DNA structure.

require Mg2+ and C02+

46
Q

DNA can be denatured by _____ and renatured through _____.

A

heating, annealing

47
Q

What does heating do to DNA’s structure and absorbance? What happens to DNA’s uv absorbance?

A

When a solution of duplex DNA is heated above a characteristic temperature, its native structure collapses and its two complementary strands separate and assume random conformations
viscosity decreases
Likewise, DNA’s ultraviolet absorbance, which is almost entirely due to its aromatic bases, increases by ∼40% on denaturation (Fig. 24-21) as a consequence of the disruption of the electronic interactions among neighboring bases. The increase in absorbance is known as the hyperchromic effect.

48
Q

What is the Tm? What does it depend on? When does it increase?

A

Tm is the melting temperature, is the temperature at which half of the maximum absorbance increase is attained.The temperature at midpoint absorbance

Tm depends on several factors including the solvent, ion in the solution, pH, and base composition.
When GC content is higher in DNA Tm increases due to their greater stacking energy

49
Q

How can Denatured DNA be renatured? What does rapid cooling or directly placing it on ice cause?

A

by cooling
Denatured DNA can be renatured when temperature is cooled below its Tm.

If temperature is maintained ~25oC below the Tm, short base-paired regions can rearrange by melting and reforming. This condition is called annealing condition.

Rapid cooling down will cause partial base pairing because the complementary strands will not have had sufficient time to find each other before the randomly base-paired structure becomes effectively “frozen in.”

50
Q

Bacterial ribosomes contain how much protein and RNA? Smallest rRNA? Describe it/What is the primary cause for single-strand RNA to form stem-and-loop structure?

A

Bacterial ribosomes contain 1/3 proteins and 2/3 RNA.

  • 5S rRNA is the smallest one containing 120 residues.
  • 5S rRNA forms several stem-and-loop structure by intramolecular Watson-Crick base pairing and non-Watson-Crick base pairing.
51
Q

tRNA Molecules Are Stabilized by?

A

Stacking Interactions.

52
Q

some RNAs are? One of the best characterized ribozymes?

A

catalysts. Ribozymes.

The hammerhead ribozyme, initially found in the RNA of certain plant virus, are RNAs that exhibit enzyme-like catalytic activity to cleave RNA during posttranscriptional RNA processing.
RNA that acts as an enzyme

53
Q

Nucleic acids can be fractionated/isolated on the basis of size, composition, and sequence by what 3 techniques?

A

Affinity Chromatography, Electrophoresis, Ultracentrifugation, and maybe solubilization

54
Q

Describe how Affinity Chromatography is used to fractionate nucleic acids?

Double-stranded DNA binds to what more tightly?

A

Double-stranded DNA binds to hydroxyapatite more tightly than do most other molecules. Therefore, hydroxyapatite column can be used to purify DNA from protein and RNA.

  • mRNA contains poly(A) tail. They can be isolated by binding to poly (T) attached to agarose or cellulose.
55
Q

Describe how Polyacrylamide Gel Electrophoresis is used to fractionate nucleic acids?
Southern, Northern, and Western blotting are used to detect what respectively?

A

Separates by size
Agarose gels can be used to resolve large fragments of DNA. Polyacrylamide gels are used to separate shorter nucleic acids
Agarose gels typically separate DNA in the double-stranded form while polyacrylamide gels separate DNA in the single-stranded form.

  • Polyacrylamide gel electrophoresis—for example, DNA sequencing.
  • Agarose gel electrophoresis—regularly used in molecular biology research.
  • Southern Blotting—detect specific DNA sequence.
  • Northern Blotting—detect specific RNA sequence; generally used to detect transcript level of genes.
  • Western Blotting—detect specific protein using antibody against the protein.

However, DNAs of more than a few thousand base pairs cannot penetrate even a weakly cross-linked polyacrylamide gel and so must be separated in agarose gels. Yet conventional gel electrophoresis is limited to DNAs of <100,000 bp, because larger DNA molecules tend to worm their way through the agarose at a rate independent of their size.

56
Q

Describe how Ultracentrifugation is used to fractionate nucleic acids?
What is the commonly used DNA separation procedure and examples?

A

Equilibrium density gradient ultracentrifugation in CsCl is a commonly used DNA separation procedure.

Equilibrium density gradient ultracentrifugation to separate DNAs
* Separate DNAs with higher G+C content from lower G+C content;
* Separate single-stranded DNA from double-stranded DNA;
* Separate circular DNA from linear DNA.

57
Q

The mechanism by which intercalating agents stain DNA

A

Intercalation Agents Stain Duplex DNA
* DNA in gels or cells can be stained by planar acromatic cations such as ethidium ion, proflavin, and acridine orange.
* These dyes bind to double-stranded DNA by intercalation (slipping in between the stacked bases), exhibiting a fluorescence under UV light that is far more intense than that of the free dye.
*Intercalating agents cause mutations in DNA and are usually carcinogens.

A substance that inserts itself into the DNA structure of a cell and binds to the DNA. This causes DNA damage.

58
Q

The accessibility of genetic information depends on the ability of proteins to?

A

recognize and interact with DNA in a manner that allows the encoded information to be copied as DNA (in replication) or as RNA (in transcription).

59
Q

What is Non-specific protein-DNA binding? examples

A

-protein can bind to any DNA sequence, for example, histone-DNA interaction, some proteins in the DNA replication machinery.
bind to DNA primarily through interactions between protein functional groups and the sugar–phosphate backbone of DNA

binds without regard to the sequence of nucleotide

60
Q

What is Sequence-specific protein DNA binding? examples

A

protein binds to specific DNA sequence, for example, restriction endonuclease binds to its recognition site in DNA, transcription factor binds to specific DNA motif. Sequence-specific DNA binding proteins presumably also bind nonspecifically but loosely to DNA so that they can scan the DNA chain for their target sequences before binding specifically and tightly.

Sequence-specific DNA-binding proteins interact primarily with bases in the major groove and with phosphate groups through direct and indirect hydrogen bonds, van der Waals interactions, and ionic interactions. The conformations of both the protein and the DNA may change on binding.

61
Q

Type II restriction does what to DNA upon binding?

A

Distort.
ype II restriction endonucleases rid bacterial cells of foreign DNA by cleaving the DNA at specific sites that have not yet been methylated by the host’s modification methylase
Upon protein binding, the DNA is distorted and major groove is widened in the recognition site.
The N-terminal ends of each subunit are inserted into the widened major groove, where they
participate in a hydrogen-bonded network with the bases of the recognition sequence.

62
Q

The purpose of structural motifs in DNA? Common structural motifs in DNA-binding proteins in prokaryotes?

A

achieves the recognition of specific DNA sequence
the helix–turn–helix (HTH) motif in prokaryotic repressors

63
Q

Common structural motifs in DNA-binding proteins in eukaryotes?

A

zinc fingers leucine zippers, and basic helix–loop–helix (bHLH) motifs in eukaryotic transcription factors.

64
Q

In prokaryotes, the expression of many genes is governed at least in part by ? What motif do they contain

A

repressors, proteins that bind at or near the gene so as to prevent its transcription.

Prokaryotic repressors interact with DNA by inserting a helix or a β strand into the major groove.

A helix-turn-helix motif within the DNA-binding protein achieves the recognition of specific DNA sequence. One α helix in this motif, called recognition helix (R), fits into the major groove of the DNA, allowing the amino acid residues to interact with the base pairs.

65
Q

What are HTF motifs? What is the recognition helix?

A

A helix-turn-helix motif contain 2 a helices
all of them bind DNA, found in proteins to regulate gene expression
binds to major groove of DNA in prokaryotes

66
Q

The E. coli trp repressor binds DNA indirectly. Describe the process

A
  • The trp operon functions to encode enzymes that are required for tryptophan synthesis.

The operator (repressor recognition sequence) assumes a sequence-specific confirmation that makes favorable contacts with the repressor rather than direct binding to the specificsequence.

67
Q

In eukaryotes, genes are selectively expressed in different cell types; this requires more complicated regulatory machinery than in prokaryotes. Prokaryotic repressors of known structure either contain an HTH motif or resemble the met repressor. However, eukaryotic DNA-binding proteins employ a much wider variety of structural motifs to bind DNA. A number of proteins known as transcription factors What are these

A

A number of proteins known as transcription factors promote the transcription of genes by binding to specific DNA sequences at or near those genes.

  1. HTH
    II. Zinc Containing DNA Binding Domains
    this motif requires a zinc molecule. It is characterized by two β sheets, a turn, and one α helix. The α helix binds to the major groove. The key thing is that you have cysteine and histidine residues that are connected by one zinc molecule
    III. Leucine Zipper Motif
    This DNA binding motif consists of Two α helical form both the dimerization and DNA-binding domain.
    embraces DNA by inserting into the major groove on opposite sides of the DNA-helix

IV. Helix-Loop-Helix (HLH) Proteins
This DNA binding motif consists consists of a short alpha chain connected by a loop to a second longer α chain. It has 3 domains: DNA binding domain, dimerization domain, activation domain.

68
Q

In terms of the way it interacts with DNA, the helix-loop-helix motif is more closely related to the leucine zipper motif why

A

Both the helix-loop -helix and the leucine zipper motif are structural motifs that allow transcription regulators to dimerize, so that each member of the pair can position an α helix in the major groove of the DNA.

69
Q

Generally transcription factors interact with the major groove or minor groove in DNA duplex?

A

major groove

70
Q

Generally transcription factors act as a dimmer or monomer?

A

dimmer

71
Q

Process of DNA packing.

A
  1. DNA wraps around histone proteins to form nucleosomes (histones are small, basic, and chromatin associated proteins.)
  2. nucleosomes further condense to form chromatin the resulting DNA-protein complex
    (Nucleosomes are the beadlike structure in eukaryotic chromatin and is the fundamental structural unit of chromatin.)
72
Q

Nucleosome structure.
How many histones?
Core DNA?
Link DNA?
DNA length in each?
purpose of the formation of nucleosomes?

A
  • Composed of a core of eight histone proteins and the DNA wrapped around them.
  • Formation of nucleosomes is the first step in compacting DNA (6-fold) into 1,000 to 10,000-fold.
  • Core DNA: the DNA most tightly associated with the nucleosome. It is wound about

1.65 times around the outside of the histone octamer.

  • Linker DNA: the DNA between each nucleosome.
    nucleosomes are joined bu the DNA between them
  • ~146 bp length of core DNA is an invariant feature of nucleosomes in all eukaryotic cells, but the liker DNA is variable (20-60bp).
73
Q

Histone structure?
Pos/Neg?
Linker histone?
is it conserved?

A
  • Histone are small, positively-charged proteins (>20% Lysine and Arginine).
  • Core histones: formed by H2A, H2B, H3 and H4 and wrapped by core DNA.
  • Linker histone: H1, binding to the linker DNA.
    brings nucleosomes together
  • N-terminal tail: different from each other and accessible within the intact nucleosome. This region is subjected to several forms of covalentmodification, e.g. acetylation, methylation and phosphorylation.
74
Q

Nucleosome Structure: Assembly

A

The assembly of a nucleosome is mediatedby the formation of H3-H4 tetramer.

Thetetramer than binds to dsDNA. The H3-H4tetramer forms a scaffold of the octamer onto which two H2A-H2B dimers are added, to complete the assembly.

75
Q
  • Eukaryotic chromatin contains how many histones?
    How about nucleosomes?
A
  • Eukaryotic chromatin contains five histones: H1, H2A, H2B, H3, H4.
    *Nucleosome has 8
76
Q

How does H1 mediates higher-order chromatin structure

A
  • Histone H1 is a small, positively-charged protein.
  • H1 is less well conserved so provides more stability
  • H1 binds to two distinct regions of DNA duplex: the linker DNA at one end of the nucleosome and the middle of the core DNA of the nucleosome.
  • H1 binding increases the length of the DNA wrapped tightly around the histone octamer by bringing these two region into close proximity, resulting in a ZigZag appearance of the nucleosomal DNA.