ELM
The Eukaryotic Linear Motif resource for
Functional Sites in Proteins

LIG_Actin_WH2_2

Accession:
Functional site class:
Actin-binding motifs
Functional site description:
The function of actin in the cell is regulated in a variety of ways, including interactions with the RPEL motifs and WH2 motifs, which bind to actin's hydrophobic cleft formed by its subdomains 1 and 3. RPEL motifs, unstructured in solution, form two helices linked by a loop upon binding. They are known to sequester MRTF transcription factors in the cytosol. RPEL-containing proteins have been found in metazoans, as well as in fungi. WH2 motifs are present in many eukaryotic organisms, including metazoa, plants and fungi. The structure of the motif comprises a short N-terminal helix of a variable length followed by an irregular region. It acts via interactions with G-actin, thus promoting or inhibiting the filament assembly. In some proteins WH2 motifs are also involved in regulation of the filament nucleation process. The differences in the functioning of various WH2-containing proteins may result from motif variants, which in several cases are not fully consistent with the assumed conservation pattern.
ELMs with same func. site: LIG_Actin_RPEL_3 LIG_Actin_WH2_1 LIG_Actin_WH2_2
ELM Description:
WH2 is a bipartite motif consisting of a short alpha-helix at its N-terminus followed by a disordered region. The most important amino acids determining the motif's ability to interact with actin are leucine and lysine (often substituted by arginine) in positions -4 and -3 from the motif C-terminus, respectively. The N-terminal helix is rich in hydrophobic amino acids enabling it to fit into the cleft formed by actin's subdomains 1 and 3. The helix contains a fragment of variable length (4-7 residues) that enables another helix turn to be included. It is followed by a loop, comprising the critical leucine and lysine/arginine residues interacting directly with the actin molecule. In several cases the C-terminal region is longer and contains residues forming a conserved pattern .T.[DE]...P which is putatively stabilizing WH2- actin interactions. This extension of the motif, not included in the ELM regular expression occurs in beta-thymosins, a group of proteins composed of WH2 domains only. For proper functioning they require an additional K/R residue at the C-terminus. They also contain a phenylalanine in position -5, conserved exclusively within this protein family. However, the level of conservation in these positions is too low to consider them an integral part of the motif. The WH2 motif is present in many proteins in various eukaryotes. It is possible to find several sequence variants, which differ from the assumed conservation pattern. In some proteins there are single cases of e.g. K/R substituted by T or N, yet their role and putative impact on actin binding remains unclear. WH2 motifs were divided into two groups. In this version positions 1 and 3 are not precisely defined, as many variants are found in different proteins. Despite lower level of conservation, this group of the WH2 motifs appears to demonstrate equally high actin-binding affinity.
Pattern: [^R]..((.[ILMVF])|([ILMVF].))[^P][^P][ILVM].{4,7}L(([KR].)|(NK))[VATIGS]
Pattern Probability: 0.0006603
Present in taxon: Eukaryota
Interaction Domain:
Actin (PF00022) Actin (Stochiometry: 1 : 1)
PDB Structure: 2D1K
o See 17 Instances for LIG_Actin_WH2_2
Actin is a core structural component of the cytoskeleton, which owing to its ability to contract, polymerize and interact with a wide range of proteins influences many cellular functions such as the control of cell shape, motility, and polarity, as well as cellular transport. Both formation of filaments, and actin's action as a molecular switch can be regulated by a variety of factors, including interactions with specific motifs, which mediate actin-protein interactions by binding to actin's hydrophobic cleft formed by subdomains 1 and 3. RPEL motifs and WH2 motifs are present in a wide range of proteins and bind actin monomers in a hydrophobic cleft. Although the invariant amino acids that are essential to binding vary (arginine in RPEL and leucine in WH2, respectively), both motifs have a preference for peripheral hydrophobic amino acids (mostly leucines and isoleucines) to stabilize the interactions.
The RPEL motif is unstructured in solution and forms two helices linked by a short loop upon actin binding. The WH2 motif forms a short helix followed by an irregular region, which stabilizes upon actin binding. In both cases the residues essential for binding lie within the unstructured loop. The function of RPEL motifs was best characterized in MRTF-A and MRTF-B proteins known to interact with serum response factor transcription factor and thus activate the transcription. There, RPEL-mediated G-actin binding (regulated by Rho-signalling controlled actin polymerization) causes the protein's retention in the cytoplasm by blocking the NLS and thus prevents the formation of the MRTF-SRF transcription- activating complex. The WH2 motif, present in a variety of proteins, functions as a regulator of actin dynamics. In beta-thymosins and WASP, binding to actin inhibits its polymerization, while in ciboulot and the WAVE1/Scar family it promotes actin filament assembly. Furthermore, in WASP the WH2 motif also prevents actin filaments from spontaneous nucleation. The WH2 motifs were also found in the Phosphatase and actin regulator protein PHAR but there their function remains unclear. Although the sequences of the motifs are distinct, they bind in the same actin region (a cleft formed by subdomains 1 and 3). RPEL- and WH2- mediated actin- binding appears to be affecting a variety of significant cellular processes and seems to be essential for proper cell functioning.
o 7 selected references:


o 6 GO-Terms:
Biological Process:
Regulation Of Arp2/3 Complex-Mediated Actin Nucleation (also annotated in these classes: LIG_Actin_WH2_1 LIG_GBD_WASP_1 )
Regulation Of Barbed-End Actin Filament Capping (also annotated in these classes: LIG_Actin_WH2_1 )
Regulation Of Actin Polymerization Or Depolymerization (also annotated in these classes: LIG_Actin_WH2_1 )
Regulation Of Actin Nucleation (also annotated in these classes: LIG_Actin_WH2_1 )
Cellular Compartment:
Cytosol (also annotated in these classes: CLV_C14_Caspase3-7 CLV_Separin_Fungi CLV_Separin_Metazoa DEG_APCC_DBOX_1 DEG_APCC_KENBOX_2 DEG_APCC_TPR_1 DEG_Cend_DCAF12_1 DEG_Cend_FEM1AC_1 DEG_Cend_FEM1B_2 DEG_Cend_KLHDC2_1 DEG_Cend_TRIM7_1 DEG_COP1_1 DEG_CRBN_cyclicCter_1 DEG_Kelch_actinfilin_1 DEG_Kelch_Keap1_1 DEG_Kelch_Keap1_2 DEG_Kelch_KLHL12_1 DEG_Kelch_KLHL3_1 DEG_MDM2_SWIB_1 DEG_Nend_Nbox_1 DEG_Nend_UBRbox_1 DEG_Nend_UBRbox_2 DEG_Nend_UBRbox_3 DEG_Nend_UBRbox_4 DEG_ODPH_VHL_1 DEG_SCF_FBW7_1 DEG_SCF_FBW7_2 DEG_SCF_FBXO31_1 DEG_SCF_SKP2-CKS1_1 DEG_SCF_TRCP1_1 DEG_SIAH_1 DOC_AGCK_PIF_1 DOC_AGCK_PIF_2 DOC_AGCK_PIF_3 DOC_ANK_TNKS_1 DOC_CDC14_PxL_1 DOC_CKS1_1 DOC_CYCLIN_D_Helix_1 DOC_CYCLIN_RevRxL_6 DOC_CYCLIN_RxL_1 DOC_CYCLIN_yClb1_LxF_4 DOC_CYCLIN_yClb3_PxF_3 DOC_CYCLIN_yCln2_LP_2 DOC_GSK3_Axin_1 DOC_MAPK_DCC_7 DOC_MAPK_FxFP_2 DOC_MAPK_gen_1 DOC_MAPK_GRA24_9 DOC_MAPK_HePTP_8 DOC_MAPK_JIP1_4 DOC_MAPK_MEF2A_6 DOC_MAPK_NFAT4_5 DOC_MAPK_RevD_3 DOC_MIT_MIM_1 DOC_PP1_MyPhoNE_1 DOC_PP1_RVXF_1 DOC_PP1_SILK_1 DOC_PP2A_B56_1 DOC_PP2A_KARD_1 DOC_PP2B_LxvP_1 DOC_PP2B_PxIxIT_1 DOC_PUB_PIM_1 DOC_RSK_DDVF_1 DOC_SPAK_OSR1_1 DOC_TBK1_STING_1 DOC_WD40_RPTOR_TOS_1 DOC_WW_Pin1_4 LIG_14-3-3_CanoR_1 LIG_14-3-3_ChREBP_3 LIG_14-3-3_CterR_2 LIG_ActinCP_CPI_1 LIG_ActinCP_TwfCPI_2 LIG_Actin_RPEL_3 LIG_Actin_WH2_1 LIG_ANK_PxLPxL_1 LIG_AP2alpha_1 LIG_AP2alpha_2 LIG_APCC_ABBA_1 LIG_APCC_Cbox_1 LIG_APCC_Cbox_2 LIG_AP_GAE_1 LIG_Arc_Nlobe_1 LIG_ARL_BART_1 LIG_BH_BH3_1 LIG_BIR_II_1 LIG_BIR_III_1 LIG_BIR_III_2 LIG_BIR_III_3 LIG_BIR_III_4 LIG_CaM_1-14-15-16_REV_1 LIG_CaM_1-26_7 LIG_CaM_1-5-10-14_3 LIG_CaM_1-8-14_4 LIG_CaM_1-8-9-10_5 LIG_CaM_1-8_REV_2 LIG_CaM_IQ_9 LIG_CaMK_CASK_1 LIG_CaM_NSCaTE_8 LIG_CAP-Gly_1 LIG_CAP-Gly_2 LIG_Clathr_ClatBox_1 LIG_Clathr_ClatBox_2 LIG_CNOT1_NIM_1 LIG_CSK_EPIYA_1 LIG_CtBP_PxDLS_1 LIG_deltaCOP1_diTrp_1 LIG_DLG_GKlike_1 LIG_Dynein_DLC8_1 LIG_EABR_CEP55_1 LIG_EF_ALG2_ABM_1 LIG_EF_ALG2_ABM_2 LIG_EH_1 LIG_eIF4E_1 LIG_eIF4E_2 LIG_EVH1_1 LIG_EVH1_2 LIG_EVH1_3 LIG_FAT_LD_1 LIG_FERM_MyoX_1 LIG_FZD_DVL_PDZ LIG_G3BP_FGDF_1 LIG_GBD_Chelix_1 LIG_GBD_WASP_1 LIG_GSK3_LRP6_1 LIG_GYF LIG_IBAR_NPY_1 LIG_IRF7_LxLS_2 LIG_IRFs_LxIS_1 LIG_KLC1_WD_1 LIG_KLC1_Yacidic_2 LIG_LIR_Apic_2 LIG_LIR_Gen_1 LIG_LIR_LC3C_4 LIG_LIR_Nem_3 LIG_LYPXL_L_2 LIG_LYPXL_S_1 LIG_LYPXL_yS_3 LIG_MYND_3 LIG_OCRL_FandH_1 LIG_PAM2_1 LIG_PAM2_2 LIG_PDZ_Class_1 LIG_PDZ_Class_2 LIG_PDZ_Class_3 LIG_PDZ_Wminus1_1 LIG_Pex14_1 LIG_Pex14_2 LIG_Pex14_3 LIG_Pex14_4 LIG_Pex3_1 LIG_PIP2_ANTH_1 LIG_PIP2_ENTH_1 LIG_PROFILIN_1 LIG_PTAP_UEV_1 LIG_PTB_Apo_2 LIG_PTB_Phospho_1 LIG_SH2_CRK LIG_SH2_GRB2like LIG_SH2_NCK_1 LIG_SH2_PTP2 LIG_SH2_SFK_2 LIG_SH2_SFK_CTail_3 LIG_SH2_STAT3 LIG_SH2_STAT5 LIG_SH2_STAT6 LIG_SH3_1 LIG_SH3_2 LIG_SH3_3 LIG_SH3_4 LIG_SH3_CIN85_PxpxPR_1 LIG_SH3_PxRPPK_7 LIG_SH3_PxxDY_5 LIG_SH3_PxxPPRxxK_8 LIG_SH3_PxxxRxxKP_6 LIG_SPRY_1 LIG_SUFU_1 LIG_SxIP_EBH_1 LIG_TPR LIG_TRAF2like_MATH_loPxQ_2 LIG_TRAF2like_MATH_shPxQ_1 LIG_TRAF3_MATH_PxP_3 LIG_TRAF4_MATH_1 LIG_TRAF6_MATH_1 LIG_TYR_ITAM LIG_TYR_ITIM LIG_TYR_ITSM LIG_UFM1_UFIM_1 LIG_VCP_SHPBox_1 LIG_VCP_VBM_3 LIG_VCP_VIM_2 LIG_Vh1_VBS_1 LIG_WH1 LIG_WRC_WIRS_1 LIG_WW_1 LIG_WW_2 LIG_WW_3 MOD_AAK1BIKe_LxxQxTG_1 MOD_CAAXbox MOD_CDC14_SPxK_1 MOD_CDK_SPK_2 MOD_CDK_SPxK_1 MOD_CDK_SPxxK_3 MOD_CK1_1 MOD_CK2_1 MOD_DYRK1A_RPxSP_1 MOD_GSK3_1 MOD_LATS_1 MOD_LOK_YxT_1 MOD_NEK2_1 MOD_NEK2_2 MOD_NMyristoyl MOD_PIKK_1 MOD_PK_1 MOD_PKA_1 MOD_PKA_2 MOD_PKB_1 MOD_PLK MOD_Plk_1 MOD_Plk_2-3 MOD_Plk_4 MOD_PRMT_GGRGG_1 MOD_ProDKin_1 MOD_SPalmitoyl_2 MOD_SPalmitoyl_4 MOD_TYR_CSK MOD_TYR_DYR ELM:old_LIG_14-3-3_1 ELM:old_LIG_14-3-3_2 ELM:old_LIG_14-3-3_3 TRG_AP2beta_CARGO_1 TRG_Cilium_Arf4_1 TRG_Cilium_RVxP_2 TRG_DiLeu_BaEn_1 TRG_DiLeu_BaEn_2 TRG_DiLeu_BaEn_3 TRG_DiLeu_BaEn_4 TRG_DiLeu_BaLyEn_6 TRG_DiLeu_LyEn_5 TRG_ENDOCYTIC_2 TRG_ER_diArg_1 TRG_ER_diLys_1 TRG_ER_FFAT_1 TRG_ER_FFAT_2 TRG_Golgi_diPhe_1 TRG_LysEnd_APsAcLL_1 TRG_LysEnd_APsAcLL_3 TRG_LysEnd_GGAAcLL_1 TRG_LysEnd_GGAAcLL_2 TRG_NES_CRM1_1 TRG_NESrev_CRM1_2 TRG_PTS1 TRG_PTS2 )
Molecular Function:
Actin Binding (also annotated in these classes: LIG_Actin_RPEL_3 LIG_Actin_WH2_1 LIG_PROFILIN_1 LIG_WRC_WIRS_1 )


o 17 Instances for LIG_Actin_WH2_2
(click table headers for sorting; Notes column: =Number of Switches, =Number of Interactions)
Acc., Gene-, NameStartEndSubsequenceLogic#Ev.OrganismNotes
B4Y9V9 bimA
B4Y9V9_BURPE
57 74 AFNAVIDQIKKGDFKLKPVG TP 3 Burkholderia pseudomallei
1
C5IZN1 VopF
C5IZN1_VIBCL
210 226 HSKLMKELINHGAKLKKVST TP 2 Vibrio cholerae
1
Q87GE5 VPA1370
Q87GE5_VIBPA
166 182 HSKLMEELLTGGRKLKKVAT TP 4 Vibrio parahaemolyticus RIMD 2210633
1
Q87GE5 VPA1370
Q87GE5_VIBPA
135 151 DHSKLMEQIRQGVKLKSATK TP 5 Vibrio parahaemolyticus RIMD 2210633
2
Q9U1K1 spir
SPIR_DROME
400 417 PYEILMGDIRAKKYQLRKVM TP 12 Drosophila melanogaster (Fruit fly)
4
Q9U1K1 spir
SPIR_DROME
432 448 AHAMILEFIRSRPPLKKASD TP 6 Drosophila melanogaster (Fruit fly)
1
Q9U1K1 spir
SPIR_DROME
370 386 FWVQVIDELRRGVRLKKSNH TP 6 Drosophila melanogaster (Fruit fly)
1
A2VCK8 TMSB4X
A2VCK8_HUMAN
3 21 DKPDMAEIEKFDKSKLKKTE TP 7 Homo sapiens (Human)
1
P20065 Tmsb4x
TYB4_MOUSE
9 27 DKPDMAEIEKFDKSKLKKTE TP 2 Mus musculus (House mouse)
1
O43312 MTSS1
MTSS1_HUMAN
728 744 QGEDMLNAIRRGVKLKKTTT TP 5 Homo sapiens (Human)
1
O97428 cib
O97428_DROME
91 109 KNQFIAGIENFDAKKLKHTE TP 6 Drosophila melanogaster (Fruit fly)
1
O97428 cib
O97428_DROME
53 71 QQSIFEGITAFNQNNLKHTE TP 6 Drosophila melanogaster (Fruit fly)
1
P18281
ACTO_ACACA
2 18 MNPELQSAIGQGAALKHAET TP 2 Acanthamoeba castellanii
1
Q6AWX6 SCAR1
SCAR1_ARATH
757 774 EAGDFLHQIRTKQFNLRRVV TP 4 Arabidopsis thaliana (Thale cress)
1
Q5XPJ6 SCAR4
SCAR4_ARATH
1106 1123 ENDSLLEIIRSKSFNLRPAD TP 3 Arabidopsis thaliana (Thale cress)
1
Q9LP46 SCAR3
SCAR3_ARATH
955 972 ETGDFLQQIRTQQFNLRPVV TP 3 Arabidopsis thaliana (Thale cress)
1
Q5XPJ9 SCAR2
SCAR2_ARATH
1336 1353 DKDSLLAQIRNKSVNLKPAV TP 1 Arabidopsis thaliana (Thale cress)
1
Please cite: ELM-the Eukaryotic Linear Motif resource-2024 update. (PMID:37962385)

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