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Protein Physics Protein Physics Protein Physics Protein Physics A. V. Finkelstein & O. B. Ptitsyn LECTURE 13 LECTURE 13 porin
19

Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

Jun 21, 2020

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Page 1: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

Protein PhysicsProtein PhysicsProtein PhysicsProtein PhysicsA V Finkelstein amp

O B PtitsynLECTURE 13LECTURE 13

porin

One Classification of Proteins1Fibrous Proteins - mostly structural proteins

- an elongated unidimensional structure

2Membrane Proteins

g

- reside within the cellular amp intracellular membrane- transport molecules in and out of the cell

3Globular Proteins - short chains packed into a compact 25-40Aring globule- closely folded structurey- water soluble proteins- Ex egg albumin hemoglobin many enzymes

Globular Proteins amp Domainsbull Short chains packed into a globule

as tightly as organic molecules in a crystalin a crystal

Primary structurey

Secondary structures bullDomain sizes 20 -692 average ~ 100 residuesbullE h f th f ld i di id lly

Super-secondary structures (motifs)

bullEach of them folds individuallybullContain a hydrophobic core built from sheets and helices connected by loop regions

structures (motifs)

DomainsDomains

Multi-domain proteins(Tertiary structure)

γ-crystallin

XX--RAYRAY One protein various One protein various

NMRNMRStructures compatible Structures compatible p p

crystallizationcrystallization p p

with one NMR experimentwith one NMR experiment

Secondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands) are) areSecondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands) are ) are the most rigid details of proteins the most rigid details of proteins

they are determined with the smallest errorsthey are determined with the smallest errors

One protein various One protein various crystallizationcrystallization

Homologous (closely Homologous (closely related) proteinsrelated) proteins

Secondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands))Secondary structures (Secondary structures (αα heliceshelices ββ strandsstrands) ) are most conserved structural elements are most conserved structural elements They form a basis of protein classificationThey form a basis of protein classification

HH HHHemoHemo--globinglobin

HemoHemo--globinglobin

H l i h i il f ldH l i h i il f ldHomologous proteins have similar foldsHomologous proteins have similar foldsTrue but trivial True but trivial NONNON t i i lt i i lNONNON--trivialtrivialMany NONMany NON--homologous proteins have similar foldshomologous proteins have similar folds

Structural Classes of Globular ProteinsDomains1 ldquopurerdquo-α - a core built exclusively from alpha-helices2 ldquopurerdquo-β - a core built exclusively from (mostly antiparallel) beta-sheets3 ldquomixedrdquo αβ mixture of ldquopurerdquo α and ldquopurerdquo β3 mixed -αβ - mixture of pure -α and pure -β4 α+β - built from combinations of β-α-β motifs

Further Simplified Models of Protein Structures

Architecture of protein

Topology of

Fold Folding Pattern Packing

Topology of protein

Level of Simplification

Usefulness classification and outlining their typical features

β-proteinsbull Mostly antiparallel arrangement bull Two sheets packed against each other

bull Basic packing types Top views

Orthogonal Packing Aligned PackingBottom views

β-barrelcylinderβ y

Retinol-binding protein

PorinPorinβ-hairpin

Meander (up amp down) motifβ-hairpin

55 55Acid protease

11

445rsquo5rsquo66332rsquo2rsquo11

44662rsquo2rsquo 33 5rsquo5rsquo

22

1122

11

Serine proteaseSerine protease

Topology Diagramsp gy g

Greek keymotif

O l i l

Greek keyOverlapping loop

uncommon

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 2: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

One Classification of Proteins1Fibrous Proteins - mostly structural proteins

- an elongated unidimensional structure

2Membrane Proteins

g

- reside within the cellular amp intracellular membrane- transport molecules in and out of the cell

3Globular Proteins - short chains packed into a compact 25-40Aring globule- closely folded structurey- water soluble proteins- Ex egg albumin hemoglobin many enzymes

Globular Proteins amp Domainsbull Short chains packed into a globule

as tightly as organic molecules in a crystalin a crystal

Primary structurey

Secondary structures bullDomain sizes 20 -692 average ~ 100 residuesbullE h f th f ld i di id lly

Super-secondary structures (motifs)

bullEach of them folds individuallybullContain a hydrophobic core built from sheets and helices connected by loop regions

structures (motifs)

DomainsDomains

Multi-domain proteins(Tertiary structure)

γ-crystallin

XX--RAYRAY One protein various One protein various

NMRNMRStructures compatible Structures compatible p p

crystallizationcrystallization p p

with one NMR experimentwith one NMR experiment

Secondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands) are) areSecondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands) are ) are the most rigid details of proteins the most rigid details of proteins

they are determined with the smallest errorsthey are determined with the smallest errors

One protein various One protein various crystallizationcrystallization

Homologous (closely Homologous (closely related) proteinsrelated) proteins

Secondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands))Secondary structures (Secondary structures (αα heliceshelices ββ strandsstrands) ) are most conserved structural elements are most conserved structural elements They form a basis of protein classificationThey form a basis of protein classification

HH HHHemoHemo--globinglobin

HemoHemo--globinglobin

H l i h i il f ldH l i h i il f ldHomologous proteins have similar foldsHomologous proteins have similar foldsTrue but trivial True but trivial NONNON t i i lt i i lNONNON--trivialtrivialMany NONMany NON--homologous proteins have similar foldshomologous proteins have similar folds

Structural Classes of Globular ProteinsDomains1 ldquopurerdquo-α - a core built exclusively from alpha-helices2 ldquopurerdquo-β - a core built exclusively from (mostly antiparallel) beta-sheets3 ldquomixedrdquo αβ mixture of ldquopurerdquo α and ldquopurerdquo β3 mixed -αβ - mixture of pure -α and pure -β4 α+β - built from combinations of β-α-β motifs

Further Simplified Models of Protein Structures

Architecture of protein

Topology of

Fold Folding Pattern Packing

Topology of protein

Level of Simplification

Usefulness classification and outlining their typical features

β-proteinsbull Mostly antiparallel arrangement bull Two sheets packed against each other

bull Basic packing types Top views

Orthogonal Packing Aligned PackingBottom views

β-barrelcylinderβ y

Retinol-binding protein

PorinPorinβ-hairpin

Meander (up amp down) motifβ-hairpin

55 55Acid protease

11

445rsquo5rsquo66332rsquo2rsquo11

44662rsquo2rsquo 33 5rsquo5rsquo

22

1122

11

Serine proteaseSerine protease

Topology Diagramsp gy g

Greek keymotif

O l i l

Greek keyOverlapping loop

uncommon

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 3: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

Globular Proteins amp Domainsbull Short chains packed into a globule

as tightly as organic molecules in a crystalin a crystal

Primary structurey

Secondary structures bullDomain sizes 20 -692 average ~ 100 residuesbullE h f th f ld i di id lly

Super-secondary structures (motifs)

bullEach of them folds individuallybullContain a hydrophobic core built from sheets and helices connected by loop regions

structures (motifs)

DomainsDomains

Multi-domain proteins(Tertiary structure)

γ-crystallin

XX--RAYRAY One protein various One protein various

NMRNMRStructures compatible Structures compatible p p

crystallizationcrystallization p p

with one NMR experimentwith one NMR experiment

Secondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands) are) areSecondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands) are ) are the most rigid details of proteins the most rigid details of proteins

they are determined with the smallest errorsthey are determined with the smallest errors

One protein various One protein various crystallizationcrystallization

Homologous (closely Homologous (closely related) proteinsrelated) proteins

Secondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands))Secondary structures (Secondary structures (αα heliceshelices ββ strandsstrands) ) are most conserved structural elements are most conserved structural elements They form a basis of protein classificationThey form a basis of protein classification

HH HHHemoHemo--globinglobin

HemoHemo--globinglobin

H l i h i il f ldH l i h i il f ldHomologous proteins have similar foldsHomologous proteins have similar foldsTrue but trivial True but trivial NONNON t i i lt i i lNONNON--trivialtrivialMany NONMany NON--homologous proteins have similar foldshomologous proteins have similar folds

Structural Classes of Globular ProteinsDomains1 ldquopurerdquo-α - a core built exclusively from alpha-helices2 ldquopurerdquo-β - a core built exclusively from (mostly antiparallel) beta-sheets3 ldquomixedrdquo αβ mixture of ldquopurerdquo α and ldquopurerdquo β3 mixed -αβ - mixture of pure -α and pure -β4 α+β - built from combinations of β-α-β motifs

Further Simplified Models of Protein Structures

Architecture of protein

Topology of

Fold Folding Pattern Packing

Topology of protein

Level of Simplification

Usefulness classification and outlining their typical features

β-proteinsbull Mostly antiparallel arrangement bull Two sheets packed against each other

bull Basic packing types Top views

Orthogonal Packing Aligned PackingBottom views

β-barrelcylinderβ y

Retinol-binding protein

PorinPorinβ-hairpin

Meander (up amp down) motifβ-hairpin

55 55Acid protease

11

445rsquo5rsquo66332rsquo2rsquo11

44662rsquo2rsquo 33 5rsquo5rsquo

22

1122

11

Serine proteaseSerine protease

Topology Diagramsp gy g

Greek keymotif

O l i l

Greek keyOverlapping loop

uncommon

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 4: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

XX--RAYRAY One protein various One protein various

NMRNMRStructures compatible Structures compatible p p

crystallizationcrystallization p p

with one NMR experimentwith one NMR experiment

Secondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands) are) areSecondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands) are ) are the most rigid details of proteins the most rigid details of proteins

they are determined with the smallest errorsthey are determined with the smallest errors

One protein various One protein various crystallizationcrystallization

Homologous (closely Homologous (closely related) proteinsrelated) proteins

Secondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands))Secondary structures (Secondary structures (αα heliceshelices ββ strandsstrands) ) are most conserved structural elements are most conserved structural elements They form a basis of protein classificationThey form a basis of protein classification

HH HHHemoHemo--globinglobin

HemoHemo--globinglobin

H l i h i il f ldH l i h i il f ldHomologous proteins have similar foldsHomologous proteins have similar foldsTrue but trivial True but trivial NONNON t i i lt i i lNONNON--trivialtrivialMany NONMany NON--homologous proteins have similar foldshomologous proteins have similar folds

Structural Classes of Globular ProteinsDomains1 ldquopurerdquo-α - a core built exclusively from alpha-helices2 ldquopurerdquo-β - a core built exclusively from (mostly antiparallel) beta-sheets3 ldquomixedrdquo αβ mixture of ldquopurerdquo α and ldquopurerdquo β3 mixed -αβ - mixture of pure -α and pure -β4 α+β - built from combinations of β-α-β motifs

Further Simplified Models of Protein Structures

Architecture of protein

Topology of

Fold Folding Pattern Packing

Topology of protein

Level of Simplification

Usefulness classification and outlining their typical features

β-proteinsbull Mostly antiparallel arrangement bull Two sheets packed against each other

bull Basic packing types Top views

Orthogonal Packing Aligned PackingBottom views

β-barrelcylinderβ y

Retinol-binding protein

PorinPorinβ-hairpin

Meander (up amp down) motifβ-hairpin

55 55Acid protease

11

445rsquo5rsquo66332rsquo2rsquo11

44662rsquo2rsquo 33 5rsquo5rsquo

22

1122

11

Serine proteaseSerine protease

Topology Diagramsp gy g

Greek keymotif

O l i l

Greek keyOverlapping loop

uncommon

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 5: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

One protein various One protein various crystallizationcrystallization

Homologous (closely Homologous (closely related) proteinsrelated) proteins

Secondary structures (Secondary structures (αα--heliceshelices ββ--strandsstrands))Secondary structures (Secondary structures (αα heliceshelices ββ strandsstrands) ) are most conserved structural elements are most conserved structural elements They form a basis of protein classificationThey form a basis of protein classification

HH HHHemoHemo--globinglobin

HemoHemo--globinglobin

H l i h i il f ldH l i h i il f ldHomologous proteins have similar foldsHomologous proteins have similar foldsTrue but trivial True but trivial NONNON t i i lt i i lNONNON--trivialtrivialMany NONMany NON--homologous proteins have similar foldshomologous proteins have similar folds

Structural Classes of Globular ProteinsDomains1 ldquopurerdquo-α - a core built exclusively from alpha-helices2 ldquopurerdquo-β - a core built exclusively from (mostly antiparallel) beta-sheets3 ldquomixedrdquo αβ mixture of ldquopurerdquo α and ldquopurerdquo β3 mixed -αβ - mixture of pure -α and pure -β4 α+β - built from combinations of β-α-β motifs

Further Simplified Models of Protein Structures

Architecture of protein

Topology of

Fold Folding Pattern Packing

Topology of protein

Level of Simplification

Usefulness classification and outlining their typical features

β-proteinsbull Mostly antiparallel arrangement bull Two sheets packed against each other

bull Basic packing types Top views

Orthogonal Packing Aligned PackingBottom views

β-barrelcylinderβ y

Retinol-binding protein

PorinPorinβ-hairpin

Meander (up amp down) motifβ-hairpin

55 55Acid protease

11

445rsquo5rsquo66332rsquo2rsquo11

44662rsquo2rsquo 33 5rsquo5rsquo

22

1122

11

Serine proteaseSerine protease

Topology Diagramsp gy g

Greek keymotif

O l i l

Greek keyOverlapping loop

uncommon

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 6: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

HH HHHemoHemo--globinglobin

HemoHemo--globinglobin

H l i h i il f ldH l i h i il f ldHomologous proteins have similar foldsHomologous proteins have similar foldsTrue but trivial True but trivial NONNON t i i lt i i lNONNON--trivialtrivialMany NONMany NON--homologous proteins have similar foldshomologous proteins have similar folds

Structural Classes of Globular ProteinsDomains1 ldquopurerdquo-α - a core built exclusively from alpha-helices2 ldquopurerdquo-β - a core built exclusively from (mostly antiparallel) beta-sheets3 ldquomixedrdquo αβ mixture of ldquopurerdquo α and ldquopurerdquo β3 mixed -αβ - mixture of pure -α and pure -β4 α+β - built from combinations of β-α-β motifs

Further Simplified Models of Protein Structures

Architecture of protein

Topology of

Fold Folding Pattern Packing

Topology of protein

Level of Simplification

Usefulness classification and outlining their typical features

β-proteinsbull Mostly antiparallel arrangement bull Two sheets packed against each other

bull Basic packing types Top views

Orthogonal Packing Aligned PackingBottom views

β-barrelcylinderβ y

Retinol-binding protein

PorinPorinβ-hairpin

Meander (up amp down) motifβ-hairpin

55 55Acid protease

11

445rsquo5rsquo66332rsquo2rsquo11

44662rsquo2rsquo 33 5rsquo5rsquo

22

1122

11

Serine proteaseSerine protease

Topology Diagramsp gy g

Greek keymotif

O l i l

Greek keyOverlapping loop

uncommon

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 7: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

Structural Classes of Globular ProteinsDomains1 ldquopurerdquo-α - a core built exclusively from alpha-helices2 ldquopurerdquo-β - a core built exclusively from (mostly antiparallel) beta-sheets3 ldquomixedrdquo αβ mixture of ldquopurerdquo α and ldquopurerdquo β3 mixed -αβ - mixture of pure -α and pure -β4 α+β - built from combinations of β-α-β motifs

Further Simplified Models of Protein Structures

Architecture of protein

Topology of

Fold Folding Pattern Packing

Topology of protein

Level of Simplification

Usefulness classification and outlining their typical features

β-proteinsbull Mostly antiparallel arrangement bull Two sheets packed against each other

bull Basic packing types Top views

Orthogonal Packing Aligned PackingBottom views

β-barrelcylinderβ y

Retinol-binding protein

PorinPorinβ-hairpin

Meander (up amp down) motifβ-hairpin

55 55Acid protease

11

445rsquo5rsquo66332rsquo2rsquo11

44662rsquo2rsquo 33 5rsquo5rsquo

22

1122

11

Serine proteaseSerine protease

Topology Diagramsp gy g

Greek keymotif

O l i l

Greek keyOverlapping loop

uncommon

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 8: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

β-proteinsbull Mostly antiparallel arrangement bull Two sheets packed against each other

bull Basic packing types Top views

Orthogonal Packing Aligned PackingBottom views

β-barrelcylinderβ y

Retinol-binding protein

PorinPorinβ-hairpin

Meander (up amp down) motifβ-hairpin

55 55Acid protease

11

445rsquo5rsquo66332rsquo2rsquo11

44662rsquo2rsquo 33 5rsquo5rsquo

22

1122

11

Serine proteaseSerine protease

Topology Diagramsp gy g

Greek keymotif

O l i l

Greek keyOverlapping loop

uncommon

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 9: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

β-barrelcylinderβ y

Retinol-binding protein

PorinPorinβ-hairpin

Meander (up amp down) motifβ-hairpin

55 55Acid protease

11

445rsquo5rsquo66332rsquo2rsquo11

44662rsquo2rsquo 33 5rsquo5rsquo

22

1122

11

Serine proteaseSerine protease

Topology Diagramsp gy g

Greek keymotif

O l i l

Greek keyOverlapping loop

uncommon

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 10: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

55 55Acid protease

11

445rsquo5rsquo66332rsquo2rsquo11

44662rsquo2rsquo 33 5rsquo5rsquo

22

1122

11

Serine proteaseSerine protease

Topology Diagramsp gy g

Greek keymotif

O l i l

Greek keyOverlapping loop

uncommon

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 11: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek kemeander greek key

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 12: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

β-sandwichβ sandwich

2255 22

44

55

66 773344 66 77Greek key 2 - 5

Greek key 3 - 6Bottom view

+ N-end11

y C-endldquocomplete Greek keyrdquo

Immunoglobulin κ

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 13: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

2211 2266 55 88 33

221166 33 88 11

2266

3388

1188

γ-crystallin β catabolite activating

protein virus coat proteinprotein

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 14: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

Greek key amp Jelly roll

Greek key

Jelly roll

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 15: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

β-proteinsβ pSo far hellip

bull Packing types orthogonal amp alignedbull Structural architecturesβ-barrel β-sandwich

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures) β-hairpin meander greek ke jell rollmeander greek key jelly roll

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 16: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

β-propeller β p p

multi-blade (45678) propeller =

three β-sandwiches

Neuraminidase

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 17: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

β-prism(helix)β p ( )bull Formed by three parallel (unusual) beta-sheets

Acyl-transferase(left-handed helix)

Pectate lyase(right-handed helix)

uncommoncommon

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 18: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

Possible topologies composed of four β-strands

Meander G k kMeander Greek keysEfimovrsquos ldquoabcd ldquo structures

Spatial structures with the same Greek key topology

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip

Page 19: Protein PhysicsProtein Physicshomes.nano.aau.dk/fp/protein-physics/Protein_Physics13.pdf · Greek key motif Olil Greek key Overlapping loop uncommon. β-pproteins ... β-pp()rism(helix)

β-proteinsβ proteins

bull Built up from mostly antiparallel β-strands (based on β-hairpins)

bull Packing types orthogonal amp alignedbull Structural architecturesSt uctu a a c tectu esβ-barrel β-sandwich β-propeller β-prism

bull Motifs present in structural architecturesbull Motifs present in structural architectures(super-secondary structures)β-hairpin meander greek key jelly rollhellip