ELM
The Eukaryotic Linear Motif resource for
Functional Sites in Proteins

LIG_PAM2_1

Accession:
Functional site class:
PAM2 motif
Functional site description:
The PABP-interacting motif 2 (PAM2) mediates binding of proteins to the MLLE/PABC peptide-binding domain found in poly(A)-binding proteins and HYD E3 ubiquitin ligases. There are two variants of the PAM2 motif that bind to overlapping but distinct sites on the MLLE domain, adopt a different orientation at their termini and consequently have different recognition elements in these regions. While the core region around a critical phenylalanine is similar in the LIG_PAM2_1 and LIG_PAM2_2 variants and makes identical contacts with the MLLE domain, the N-terminal sequence in the LIG_PAM2_1 provides a hydrophobic residue that is essential for efficient binding and is lacking in LIG_PAM2_2 motifs, while the C-terminal part of the LIG_PAM2_2 variant makes a turn and follows an alternative path on the MLLE surface that depends on an aromatic residue that is absent from LIG_PAM2_1 motifs.
ELMs with same func. site: LIG_PAM2_1 LIG_PAM2_2
ELM Description:
Binding of the LIG_PAM2_1 mainly involves hydrophobic interactions, the most important of which are mediated by the residues in position 3 (lacking in LIG_PAM2_2) and 10 (corresponds to position 5 in LIG_PAM2_2). The residues in positions 1 and 2 of the motif, most often polar or charged residues, do not participate in binding, except for eRF3, which contains two overlapping motifs, where phenylalanine in position 1 of the C-terminal motif bends back to bind in the hydrophobic pocket that is normally occupied by the hydrophobic residue at position 3, which is most frequently leucine, but in some cases phenylalanine or proline. The side chain of the asparagine or serine residue in position 4 is involved in an intermolecular salt bridge and forms an intramolecular hydrogen bond with the amide of the residue in position 6. The latter interaction stabilizes the beta-turn conformation of the peptide. The regular occurrence of proline in position 5 might be due to its propensity to form such a beta-turn. Position 7 is invariantly alanine, which is involved in different hydrophobic interactions with multiple MLLE residues. Introduction of a bulky residue at this position was found to decrease the affinity for the peptide. Position 10 is the single most important residue for binding and in most cases it is occupied by phenylalanine, but exceptionally by tryptophan in LARP4 and LARP4B, or possibly also by tyrosine in the Arabidopsis thaliana proteins CID5 and CID6. Since CID5 is the only example of an experimentally validated motif with a tyrosine in position 10, this feature might be specific for plants. The residue in position 10 occupies a hydrophobic pocket between helices 2 and 3 and is the major binding determinant. In case of a phenylalanine in this position, a hydrophobic residue is found in position 12, with a clear preference for proline. AtCID5 and AtCID6 also contain a proline at position 12. For Fungi and some TAXON:7215 sequences, the regular expression is less strict.
Pattern: ..[LFP][NS][PIVTAFL].A..(([FY].[PYLF])|(W..)).
Pattern Probability: 0.0000100
Present in taxon: Eukaryota
Interaction Domain:
PABP (PF00658) Poly-adenylate binding protein, unique domain (Stochiometry: 1 : 1)
PDB Structure: 3KUS
o See 22 Instances for LIG_PAM2_1
The PABP-interacting motif 2 (PAM2) directly binds to the MLLE/PABC domain that is found in poly(A)-binding proteins (PABP) like PABPC1 and in members of the HECT domain-containing Hyperplastic Discs (HYD) protein family of E3 ubiquitin ligases (Albrecht,2004, Lim,2006). This domain consists of a conserved bundle of five alpha-helices, of which the N-terminal helix is lacking in some proteins. The interaction with PAM2 sequences involves the most conserved helices 2, 3 and 5, with the exception of yeast, where only helices 2 and 3 mediate binding.
PABPC1 binds to the 3’-poly(A) tail of mRNA molecules via four RNA recognition motifs (RRMs) and recruits different regulatory proteins, which modulate translational activity and mRNA stability, by binding to their PAM2 motifs via its C-terminal MLLE domain (Siddiqui,2007). LIG_PAM2_1 motif-containing PABPC1-binding proteins include Paip1 and Paip2, which stimulate or repress translation by stabilizing or destabilizing, respectively, the closed loop structure of mRNA that is formed by the interaction between PABP and the translation initiation factor eIF4G. Other examples are the eukaryotic release factor eRF3, which contains two overlapping LIG_PAM2_1 motifs, and Tob1 and Pan3, which recruit the deadenylase complexes Caf1-Ccr4 and Pan2-Pan3, respectively, to the mRNA poly(A) tails. These complexes are involved in translation-dependent, eRF3-mediated mRNA decay and translation termination. Tob1 and its family member Tob2 contain two distinct LIG_PAM2_1 motifs, and evidence indicates that in both cases the C-terminal motif is the main interaction site for binding to PABP. PABPC1 binding partners containing the LIG_PAM2_2 motif variant are so far restricted to animal-specific GW182 family proteins, which are part of the RNA-induced silencing complex (RISC) and are essential for microRNA (miRNA)-mediated gene silencing (Jinek,2010). Silencing is achieved by inhibiting translation and promoting degradation of mRNA, and depends on binding of GW182 proteins in the miRNA-loaded RISC to PABPC1. Additional, possibly indirect, interactions also occur between the RRMs of PABPC1 and the C-terminal region of GW182 family members. While in human GW182 proteins PABPC1 binding is predominantly mediated by their PAM2 sequences, in D. melanogaster GW182 the motif, which binds with lower affinity, also contributes to but is dispensable for this interaction in immunoprecipitation experiments (Huntzinger,2010).
The tumor suppressor protein HYD (EDD/UBR5) belongs to the family of HECT domain-containing E3 ubiquitin ligases, which target specific proteins for proteasome-dependent degradation. Evidence suggests that HYD is involved in proliferation and DNA damage signaling. Little is known about the function of the MLLE-PAM2 interaction in HYD E3 ligases, but it has been suggested that PAM2-containing proteins may be targeted for ubiquitination by HYD through binding to its MLLE domain, which is positioned directly adjacent to the catalytic HECT domain. So far, all known PAM2-mediated interactions with HYD involve instances of the LIG_PAM2_1 variant. The structural features of PAM2 binding to the MLLE domain of PABP or HYD are very similar, and similar specificity and binding affinity has been demonstrated for various peptides (Lim,2006).
o 10 selected references:


o 6 GO-Terms:
Biological Process:
Nuclear-Transcribed Mrna Catabolic Process, Deadenylation-Dependent Decay (also annotated in these classes: LIG_PAM2_2 )
Positive Regulation Of Translation (also annotated in class: )
Negative Regulation Of Translation (also annotated in these classes: LIG_CNOT1_NIM_1 LIG_PAM2_2 )
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_Actin_WH2_2 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_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:
Protein Domain Specific Binding (also annotated in these classes: DEG_APCC_DBOX_1 DEG_APCC_KENBOX_2 DEG_Cend_DCAF12_1 DEG_Cend_FEM1AC_1 DEG_Cend_FEM1B_2 DEG_Cend_KLHDC2_1 DEG_Cend_TRIM7_1 DEG_Kelch_actinfilin_1 DEG_Kelch_Keap1_1 DEG_Kelch_Keap1_2 DEG_Kelch_KLHL3_1 DOC_MIT_MIM_1 LIG_14-3-3_CanoR_1 LIG_14-3-3_CterR_2 LIG_AP_GAE_1 LIG_ARS2_EDGEI_1 LIG_BRCT_BRCA1_1 LIG_BRCT_BRCA1_2 LIG_BRCT_MDC1_1 LIG_CAP-Gly_1 LIG_CSK_EPIYA_1 LIG_CSL_BTD_1 LIG_deltaCOP1_diTrp_1 LIG_EH_1 LIG_EVH1_1 LIG_EVH1_2 LIG_EVH1_3 LIG_FHA_1 LIG_FHA_2 LIG_G3BP_FGDF_1 LIG_MSH2_SHIPbox_1 LIG_MTR4_AIM_1 LIG_PAM2_2 LIG_PDZ_Class_1 LIG_PDZ_Class_2 LIG_PDZ_Class_3 LIG_PDZ_Wminus1_1 LIG_PTB_Apo_2 LIG_PTB_Phospho_1 LIG_RRM_PRI_1 LIG_SH3_1 LIG_SH3_2 LIG_SH3_3 LIG_SH3_4 LIG_SH3_PxRPPK_7 LIG_SH3_PxxDY_5 LIG_SH3_PxxPPRxxK_8 LIG_SH3_PxxxRxxKP_6 LIG_Trf4_IWRxY_1 LIG_TRFH_1 LIG_ULM_U2AF65_1 LIG_WH1 ELM:old_LIG_14-3-3_1 ELM:old_LIG_14-3-3_2 ELM:old_LIG_14-3-3_3 )
Protein Ubiquitination (also annotated in these classes: DEG_CRBN_cyclicCter_1 DEG_SPOP_SBC_1 DOC_PUB_PIM_1 LIG_BIR_II_1 LIG_BIR_III_1 LIG_BIR_III_2 LIG_BIR_III_3 LIG_BIR_III_4 )


o 22 Instances for LIG_PAM2_1
(click table headers for sorting; Notes column: =Number of Switches, =Number of Interactions)
Acc., Gene-, NameStartEndSubsequenceLogic#Ev.OrganismNotes
Q14106 TOB2
TOB2_HUMAN
131 143 EIKSSFNPDAQVFVPIGSQD TP 8 Homo sapiens (Human)
2
Q14106 TOB2
TOB2_HUMAN
251 263 APQSQLSPNAKEFVYNGGGS TP 8 Homo sapiens (Human)
2
P50616 TOB1
TOB1_HUMAN
265 277 QKTSALSPNAKEFIFPNMQG TP 9 Homo sapiens (Human)
1
P50616 TOB1
TOB1_HUMAN
130 142 EIKNSFNPEAQVFMPISDPA TP 7 Homo sapiens (Human)
1
Q9VG13 Paip2
Q9VG13_DROME
103 115 VEKSVLNPMADEFVPRCHVI TP 4 Drosophila melanogaster (Fruit fly)
1
Q39096 ERD15
ERD15_ARATH
9 21 GRRSTLNPDAPLFIPAAVRQ TP 2 Arabidopsis thaliana (Thale cress)
1
Q6ELF8 Poly(A)-bindi
Q6ELF8_CUCSA
10 22 EGSSKLNPNAPLFIPAAYQV TP 1 Cucumis sativus (Cucumber)
1
Q6ELF9 Poly(A)-bindi
Q6ELF9_CUCSA
14 26 SSVSMLNPNAPLFVPMAYRT TP 2 Cucumis sativus (Cucumber)
1
Q92615 LARP4B
LAR4B_HUMAN
54 66 TKVSELNPNAEVWGAPVLHL TP 3 Homo sapiens (Human)
1
Q71RC2 LARP4
LARP4_HUMAN
13 25 SKGTGLNPNAKVWQEIAPGN TP 3 Homo sapiens (Human)
1
P23226 Map205
MA205_DROME
234 246 ENHSQLNPNAVAFVPGVGSQ TP 3 Drosophila melanogaster (Fruit fly)
2
P53804 TTC3
TTC3_HUMAN
1205 1217 RVKLQLNPAAREFKPDVKSK TP 2 Homo sapiens (Human)
2
Q14694 USP10
UBP10_HUMAN
81 93 SISSTLNPQAPEFILGCTAS TP 2 Homo sapiens (Human)
2
Q12986 NFX1
NFX1_HUMAN
11 23 SGTFKFNTDAAEFIPQEKKN TP 12 Homo sapiens (Human)
2
Q58A45 PAN3
PAN3_HUMAN
284 296 NNLQTPNPTASEFIPKGGST TP 7 Homo sapiens (Human)
1
Q8SWR8 Atx2
ATX2_DROME
857 869 VKKHVLNPSAKPFTPRGPST TP 2 Drosophila melanogaster (Fruit fly)
1
Q99700 ATXN2
ATX2_HUMAN
912 924 VRKSTLNPNAKEFNPRSFSQ TP 5 Homo sapiens (Human)
2
Q8IYD1 GSPT2
ERF3B_HUMAN
59 71 AKPFVPNVHAAEFVPSFLRG TP 7 Homo sapiens (Human)
2
Q8IYD1 GSPT2
ERF3B_HUMAN
50 62 AFSRKLNVNAKPFVPNVHAA TP 7 Homo sapiens (Human)
2
Q9H074 PAIP1
PAIP1_HUMAN
126 138 VLMSKLSVNAPEFYPSGYSS TP 13 Homo sapiens (Human)
2
Q9ULR5 PAIP2B
PAI2B_HUMAN
108 120 ILSKSNLNPDAKEFIPGEKY TP 5 Homo sapiens (Human)
1
Q9BPZ3 PAIP2
PAIP2_HUMAN
109 121 VVKSNLNPNAKEFVPGVKYG TP 12 Homo sapiens (Human)
2
Please cite: ELM-the Eukaryotic Linear Motif resource-2024 update. (PMID:37962385)

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