PROFILPELAJAR.COM

Available protein structures:
Pfam	structures / ECOD
PDB	RCSB PDB; PDBe; PDBj
PDBsum	structure summary
PDB	1b3u, 1f59, 1gcj, 1ibr, 1m5n, 1o6o, 1o6p, 1qbk, 1qgr, 1u6g, 1ukl, 2bku

HEAT repeat
HEAT repeat
	An example of an alpha solenoid structure composed of 15 HEAT repeats. The protein phosphatase 2A regulatory subunit is shown with the N-terminus in blue at bottom and the C-terminus in red at top. A single helix-turn-helix motif is shown in the center with the outer helix in pink, the inner helix in green, and the turn in white. From PDB: 2IAE.
Identifiers
Symbol	HEAT
Pfam	PF02985
InterPro	IPR000357
PROSITE	PDOC50077
SCOP2	1b3u / SCOPe / SUPFAM
Pfam
Available protein structures:
Pfam	structures / ECOD
PDB	RCSB PDB; PDBe; PDBj
PDBsum	structure summary
PDB	1b3u, 1f59, 1gcj, 1ibr, 1m5n, 1o6o, 1o6p, 1qbk, 1qgr, 1u6g, 1ukl, 2bku

A HEAT repeat is a protein tandem repeat structural motif composed of two alpha helices linked by a short loop. HEAT repeats can form alpha solenoids, a type of solenoid protein domain found in a number of cytoplasmic proteins. The name "HEAT" is an acronym for four proteins in which this repeat structure is found: Huntingtin, elongation factor 3 (EF3), protein phosphatase 2A (PP2A),^[3] and the yeast kinase TOR1.^[4] HEAT repeats form extended superhelical structures which are often involved in intracellular transport; they are structurally related to armadillo repeats. The nuclear transport protein importin beta contains 19 HEAT repeats.

Various HEAT repeat proteins and their structures

Representative examples of HEAT repeat proteins include importin β (also known as karyopherin β) family,^[5] regulatory subunits of condensin and cohesin,^[6] separase,^[7] PIKKs (phosphatidylinositol 3-kinase-related protein kinases) such as ATM (Ataxia telangiectasia mutated) and ATR (Ataxia telangiectasia and Rad3 related),^[8]^[9] and the microtubule-binding protein XMAP215/Dis1/TOG^[10] and CLASP.^[11] Thus, cellular functions of HEAT repeat proteins are highly variable.

The structure of the following HEAT repeat proteins have been determined so far:

Protein modification and degradation
- A subunit^[2] and holoenzyme^[12]^[13] of PP2A
- SCFubiquitin ligase regulator Cand1^[14]
- Hsm3, a molecular chaperon involved in the assembly of 26Sproteasome^[15]
Nucleo-cytoplasmic transport
- Importin β^[16]^[17]^[18]
- Exportin Cse1^[19]
- Transportin 1^[20]
- Nuleoporin Gle1;^[21] Nup188;^[22] Nup192^[23]
Transcriptional regulation
- TFIID subunit TAF6^[24]
- TBP regulator Mot1 (Modifier of transcription 1)^[25]
- Transcriptional initiation factor Rrn3^[26]
Translational regulation
- Elongation factor eEF3^[27]
- Initiation factor eIF4G^[28]
- Aminoacyl tRNA synthetase transfer protein Cex1p^[29]
DNA repair
- DNA-PK (DNA-dependent protein kinase)^[30]
- Fanconi anemia responsible protein FANCF (FANCF)^[31]
- Damaged DNA-binding protein AlkD (Alkylpurin DNA glycosylase)^[32]
- PIKKs chaperone Tel2^[33]
Chromosomal regulation
- Cohesin subunit SA2^[34]/Scc3^[35]^[36]
- Cohesin regulator Wapl^[37]^[38]
- Cohesin regulator Pds5^[39]^[40]^[41]
- Cohesin loading factor Scc2^[42]^[43]
- Cohesin protease Separase^[44]^[45]^[46]
- Condensin subunit CAP-G/ycg1^[47]^[48]
- Condensin subunit CAP-D2/ycs4^[49]
Cytoskeletal regulation
- Microtubule-binding protein TOG^[50]^[51]/Stu2^[52]
Cell proliferation regulation
- TOR (target of rapamycin)^[53]
Others
- API5 (Apoptosis inhibitor 5)^[54]
- V-type ATPase H subunit^[55]

References

^ Cho, Uhn Soo; Xu, Wenqing (1 November 2006). "Crystal structure of a protein phosphatase 2A heterotrimeric holoenzyme". Nature. 445 (7123): 53–57. doi:10.1038/nature05351. PMID 17086192. S2CID 4408160.
^ ^a ^b Groves MR, Hanlon N, Turowski P, Hemmings BA, Barford D (January 1999). "The structure of the protein phosphatase 2A PR65/A subunit reveals the conformation of its 15 tandemly repeated HEAT motifs". Cell. 96 (1): 99–110. doi:10.1016/S0092-8674(00)80963-0. PMID 9989501. S2CID 14465060.
^ Kobe, Bostjan; Gleichmann, Thomas; Horne, James; Jennings, Ian G.; Scotney, Pierre D.; Teh, Trazel (1999-05-05). "Turn up the HEAT". Structure. 7 (5): R91–R97. doi:10.1016/S0969-2126(99)80060-4. ISSN 0969-2126. PMID 10378263.
^ Andrade MA, Bork P (October 1995). "HEAT repeats in the Huntington's disease protein". Nat. Genet. 11 (2): 115–6. doi:10.1038/ng1095-115. PMID 7550332. S2CID 6911746.
^ Malik HS, Eickbush TH, Goldfarb DS (1997). "Evolutionary specialization of the nuclear targeting apparatus". Proc. Natl. Acad. Sci. USA. 94 (25): 13738–13742. Bibcode:1997PNAS...9413738M. doi:10.1073/pnas.94.25.13738. PMC 28376. PMID 9391096.{{cite journal}}: CS1 maint: multiple names: authors list (link)
^ Neuwald AF, Hirano T (2000). "HEAT repeats associated with condensins, cohesins, and other complexes involved in chromosome-related functions". Genome Res. 10 (10): 1445–52. doi:10.1101/gr.147400. PMC 310966. PMID 11042144.
^ Jäger H, Herzig B, Herzig A, Sticht H, Lehner CF, Heidmann S (2004). "Structure predictions and interaction studies indicate homology of separase N-terminal regulatory domains and Drosophila THR". Cell Cycle. 3 (2): 182–188. doi:10.4161/cc.3.2.605. PMID 14712087.{{cite journal}}: CS1 maint: multiple names: authors list (link)
^ Perry J, Kleckner N (2003). "The ATRs, ATMs, and TORs are giant HEAT repeat proteins". Cell. 112 (2): 151–155. doi:10.1016/s0092-8674(03)00033-3. PMID 12553904. S2CID 17261901.
^ Baretić D, Williams RL (2014). "PIKKs--the solenoid nest where partners and kinases meet". Curr. Opin. Struct. Biol. 29: 134–142. doi:10.1016/j.sbi.2014.11.003. PMID 25460276.
^ Ohkura, Hiroyuki; Garcia, Miguel A.; Toda, Takashi (1 November 2001). "Dis1/TOG universal microtubule adaptors - one MAP for all?". Journal of Cell Science. 114 (21): 3805–3812. doi:10.1242/jcs.114.21.3805. PMID 11719547.
^ Al-Bassam J, Kim H, Brouhard G, van Oijen A, Harrison SC, Chang F (2010). "CLASP promotes microtubule rescue by recruiting tubulin dimers to the microtubule". Dev. Cell. 19 (2): 245–258. doi:10.1016/j.devcel.2010.07.016. PMC 3156696. PMID 20708587.{{cite journal}}: CS1 maint: multiple names: authors list (link)
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^ Muir KW, Kschonsak M, Li Y, Metz J, Haering CH, Panne D. (2016). "Structure of the Pds5-Scc1 complex and implications for cohesin function". Cell Rep. 14 (9): 2116–2126. doi:10.1016/j.celrep.2016.01.078. PMID 26923589.{{cite journal}}: CS1 maint: multiple names: authors list (link)
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^ Ouyang Z, Zheng G, Tomchick DR, Luo X, Yu H. (2016). "Structural basis and IP6 requirement for Pds5-dependent cohesin dynamics". Mol Cell. 62 (2): 248–259. doi:10.1016/j.molcel.2016.02.033. PMC 5560056. PMID 26971492.{{cite journal}}: CS1 maint: multiple names: authors list (link)
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External links

Eukaryotic Linear Motif resource motif class LIG_PIKK_1
Eukaryotic Linear Motif resource motif class DOC_PIKK_1

[cho_2006-1] Cho, Uhn Soo; Xu, Wenqing (1 November 2006). "Crystal structure of a protein phosphatase 2A heterotrimeric holoenzyme". Nature. 445 (7123): 53–57. doi:10.1038/nature05351. PMID 17086192. S2CID 4408160.

[pmid9989501-2] Groves MR, Hanlon N, Turowski P, Hemmings BA, Barford D (January 1999). "The structure of the protein phosphatase 2A PR65/A subunit reveals the conformation of its 15 tandemly repeated HEAT motifs". Cell. 96 (1): 99–110. doi:10.1016/S0092-8674(00)80963-0. PMID 9989501. S2CID 14465060.

[3] Kobe, Bostjan; Gleichmann, Thomas; Horne, James; Jennings, Ian G.; Scotney, Pierre D.; Teh, Trazel (1999-05-05). "Turn up the HEAT". Structure. 7 (5): R91–R97. doi:10.1016/S0969-2126(99)80060-4. ISSN 0969-2126. PMID 10378263.

[pmid7550332-4] Andrade MA, Bork P (October 1995). "HEAT repeats in the Huntington's disease protein". Nat. Genet. 11 (2): 115–6. doi:10.1038/ng1095-115. PMID 7550332. S2CID 6911746.

[pmid9391096-5] Malik HS, Eickbush TH, Goldfarb DS (1997). "Evolutionary specialization of the nuclear targeting apparatus". Proc. Natl. Acad. Sci. USA. 94 (25): 13738–13742. Bibcode:1997PNAS...9413738M. doi:10.1073/pnas.94.25.13738. PMC 28376. PMID 9391096.{{cite journal}}: CS1 maint: multiple names: authors list (link)

[pmid11042144-6] Neuwald AF, Hirano T (2000). "HEAT repeats associated with condensins, cohesins, and other complexes involved in chromosome-related functions". Genome Res. 10 (10): 1445–52. doi:10.1101/gr.147400. PMC 310966. PMID 11042144.

[pmid14712087-7] Jäger H, Herzig B, Herzig A, Sticht H, Lehner CF, Heidmann S (2004). "Structure predictions and interaction studies indicate homology of separase N-terminal regulatory domains and Drosophila THR". Cell Cycle. 3 (2): 182–188. doi:10.4161/cc.3.2.605. PMID 14712087.{{cite journal}}: CS1 maint: multiple names: authors list (link)

[pmid12553904-8] Perry J, Kleckner N (2003). "The ATRs, ATMs, and TORs are giant HEAT repeat proteins". Cell. 112 (2): 151–155. doi:10.1016/s0092-8674(03)00033-3. PMID 12553904. S2CID 17261901.

[pmid25460276-9] Baretić D, Williams RL (2014). "PIKKs--the solenoid nest where partners and kinases meet". Curr. Opin. Struct. Biol. 29: 134–142. doi:10.1016/j.sbi.2014.11.003. PMID 25460276.

[pmid11719547-10] Ohkura, Hiroyuki; Garcia, Miguel A.; Toda, Takashi (1 November 2001). "Dis1/TOG universal microtubule adaptors - one MAP for all?". Journal of Cell Science. 114 (21): 3805–3812. doi:10.1242/jcs.114.21.3805. PMID 11719547.

[pmid20708587-11] Al-Bassam J, Kim H, Brouhard G, van Oijen A, Harrison SC, Chang F (2010). "CLASP promotes microtubule rescue by recruiting tubulin dimers to the microtubule". Dev. Cell. 19 (2): 245–258. doi:10.1016/j.devcel.2010.07.016. PMC 3156696. PMID 20708587.{{cite journal}}: CS1 maint: multiple names: authors list (link)

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HEAT repeat

Various HEAT repeat proteins and their structures

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