Skip Header

You are using a version of browser that may not display all the features of this website. Please consider upgrading your browser.
Protein

Arf-GAP with Rho-GAP domain, ANK repeat and PH domain-containing protein 3

Gene

Arap3

Organism
Mus musculus (Mouse)
Status
Reviewed-Annotation score: Annotation score: 5 out of 5-Experimental evidence at protein leveli

Functioni

Phosphatidylinositol 3,4,5-trisphosphate-dependent GTPase-activating protein that modulates actin cytoskeleton remodeling by regulating ARF and RHO family members. Is activated by phosphatidylinositol 3,4,5-trisphosphate (PtdIns(3,4,5)P3) binding. Can be activated by phosphatidylinositol 3,4-bisphosphate (PtdIns(3,4,5)P2) binding, albeit with lower efficiency. Acts preferentially on ARF5 and on RHOA.1 Publication

GO - Molecular functioni

GO - Biological processi

  • negative regulation of cell migration Source: MGI
  • negative regulation of Rac protein signal transduction Source: MGI
  • negative regulation of Rho protein signal transduction Source: MGI
  • positive regulation of GTPase activity Source: GOC
  • regulation of cell shape Source: MGI
  • signal transduction Source: InterPro
Complete GO annotation...

Keywords - Molecular functioni

GTPase activation

Enzyme and pathway databases

ReactomeiR-MMU-194840. Rho GTPase cycle.

Names & Taxonomyi

Protein namesi
Recommended name:
Arf-GAP with Rho-GAP domain, ANK repeat and PH domain-containing protein 3
Alternative name(s):
Centaurin-delta-3
Short name:
Cnt-d3
Dual specificity Rho- and Arf-GTPase-activating protein 1
Gene namesi
Name:Arap3
Synonyms:Centd3, Drag1, Kiaa4097
OrganismiMus musculus (Mouse)
Taxonomic identifieri10090 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresGliresRodentiaSciurognathiMuroideaMuridaeMurinaeMusMus
Proteomesi
  • UP000000589 Componenti: Chromosome 18

Organism-specific databases

MGIiMGI:2147274. Arap3.

Subcellular locationi

GO - Cellular componenti

  • cytoplasm Source: MGI
  • cytoskeleton Source: UniProtKB-SubCell
  • lamellipodium Source: MGI
  • plasma membrane Source: UniProtKB-SubCell
  • ruffle Source: MGI
Complete GO annotation...

Keywords - Cellular componenti

Cell membrane, Cell projection, Cytoplasm, Cytoskeleton, Membrane

Pathology & Biotechi

Mutagenesis

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Mutagenesisi938 – 9381R → L: Loss of Rho GAP activity. Enhances cell spreading. 1 Publication
Mutagenesisi1399 – 13991Y → F: Strongly reduces phosphorylation by CSK; when associated with F-1404. 1 Publication
Mutagenesisi1404 – 14041Y → F: Strongly reduces phosphorylation by CSK; when associated with F-1399. 1 Publication

PTM / Processingi

Molecule processing

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Chaini1 – 15381538Arf-GAP with Rho-GAP domain, ANK repeat and PH domain-containing protein 3PRO_0000074216Add
BLAST

Amino acid modifications

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Modified residuei1344 – 13441PhosphothreonineCombined sources
Modified residuei1399 – 13991Phosphotyrosine1 Publication
Modified residuei1404 – 14041PhosphotyrosineCombined sources1 Publication
Modified residuei1438 – 14381PhosphoserineBy similarity
Modified residuei1474 – 14741PhosphoserineBy similarity

Post-translational modificationi

Tyrosine phosphorylated at a low basal level. PDGF treatment stimulates phosphorylation. Tyrosine phosphorylation is increased in cells that are in the process of becoming attached to a substrate and that start spreading and flattening.1 Publication

Keywords - PTMi

Phosphoprotein

Proteomic databases

MaxQBiQ8R5G7.
PaxDbiQ8R5G7.
PRIDEiQ8R5G7.

PTM databases

iPTMnetiQ8R5G7.
PhosphoSiteiQ8R5G7.

Expressioni

Gene expression databases

BgeeiQ8R5G7.
GenevisibleiQ8R5G7. MM.

Interactioni

Subunit structurei

Interacts (via SAM domain) with INPPL1/SHIP2.By similarity

Binary interactionsi

WithEntry#Exp.IntActNotes
LYNP079482EBI-621463,EBI-79452From a different organism.
SRCP129313EBI-621463,EBI-621482From a different organism.

Protein-protein interaction databases

IntActiQ8R5G7. 3 interactions.
STRINGi10090.ENSMUSP00000035662.

Structurei

3D structure databases

ProteinModelPortaliQ8R5G7.
SMRiQ8R5G7. Positions 1-80.
ModBaseiSearch...
MobiDBiSearch...

Family & Domainsi

Domains and Repeats

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Domaini4 – 6865SAMPROSITE-ProRule annotationAdd
BLAST
Domaini282 – 37493PH 1PROSITE-ProRule annotationAdd
BLAST
Domaini389 – 47890PH 2PROSITE-ProRule annotationAdd
BLAST
Domaini479 – 606128Arf-GAPPROSITE-ProRule annotationAdd
BLAST
Domaini671 – 785115PH 3PROSITE-ProRule annotationAdd
BLAST
Domaini795 – 901107PH 4PROSITE-ProRule annotationAdd
BLAST
Domaini903 – 1084182Rho-GAPPROSITE-ProRule annotationAdd
BLAST
Domaini1113 – 120694Ras-associatingPROSITE-ProRule annotationAdd
BLAST
Domaini1219 – 1321103PH 5PROSITE-ProRule annotationAdd
BLAST

Compositional bias

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Compositional biasi1493 – 153644Pro-richAdd
BLAST

Sequence similaritiesi

Contains 1 Arf-GAP domain.PROSITE-ProRule annotation
Contains 5 PH domains.PROSITE-ProRule annotation
Contains 1 Ras-associating domain.PROSITE-ProRule annotation
Contains 1 Rho-GAP domain.PROSITE-ProRule annotation
Contains 1 SAM (sterile alpha motif) domain.PROSITE-ProRule annotation

Keywords - Domaini

Repeat

Phylogenomic databases

eggNOGiKOG1117. Eukaryota.
COG5347. LUCA.
GeneTreeiENSGT00760000118874.
HOGENOMiHOG000246988.
HOVERGENiHBG069114.
InParanoidiQ8R5G7.
KOiK12490.
OMAiHEGIYRK.
OrthoDBiEOG7F24T4.
TreeFamiTF105769.

Family and domain databases

Gene3Di1.10.150.50. 1 hit.
1.10.555.10. 1 hit.
2.30.29.30. 3 hits.
InterProiIPR001164. ArfGAP.
IPR011993. PH_dom-like.
IPR001849. PH_domain.
IPR000159. RA_dom.
IPR008936. Rho_GTPase_activation_prot.
IPR000198. RhoGAP_dom.
IPR001660. SAM.
IPR013761. SAM/pointed.
IPR029071. Ubiquitin-rel_dom.
[Graphical view]
PfamiPF01412. ArfGap. 1 hit.
PF00169. PH. 2 hits.
PF00788. RA. 1 hit.
PF00620. RhoGAP. 1 hit.
PF07647. SAM_2. 1 hit.
[Graphical view]
PRINTSiPR00405. REVINTRACTNG.
SMARTiSM00105. ArfGap. 1 hit.
SM00233. PH. 5 hits.
SM00324. RhoGAP. 1 hit.
SM00454. SAM. 1 hit.
[Graphical view]
SUPFAMiSSF47769. SSF47769. 1 hit.
SSF48350. SSF48350. 1 hit.
SSF50729. SSF50729. 5 hits.
SSF54236. SSF54236. 1 hit.
PROSITEiPS50115. ARFGAP. 1 hit.
PS50003. PH_DOMAIN. 3 hits.
PS50200. RA. 1 hit.
PS50238. RHOGAP. 1 hit.
PS50105. SAM_DOMAIN. 1 hit.
[Graphical view]

Sequences (3)i

Sequence statusi: Complete.

This entry describes 3 isoformsi produced by alternative splicing. AlignAdd to basket

Isoform 1 (identifier: Q8R5G7-1) [UniParc]FASTAAdd to basket

This isoform has been chosen as the 'canonical' sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

« Hide

        10         20         30         40         50
MAAPQDLDIA VWLALVHLEQ YADTFRRHGL ATAGAAQHLG HEELRHLGIS
60 70 80 90 100
ATGHRKRILR LLRAGSAEGF LDSHLDNTME PTPSPAPDAQ PPKPVPKPRT
110 120 130 140 150
VFGLSNPATA QRPGLSPIFW DPEVSRNSEC TQRSSPLLPS SSEQPSVPNT
160 170 180 190 200
MEMMPNAIYF GLDLRGRAQA AQDVTPDSSQ ATVPTPAFRP TTGTVHIMDP
210 220 230 240 250
GCLYYGVQPV GIPGASDRRD GRGVCQERAE HRQDLETRED AGYASLELPG
260 270 280 290 300
DSILSLPTQD AETSDDLISP YASFSSTADR PVPLLSGWLD KLSPQGNYVF
310 320 330 340 350
QRRFVQFNGR SLMYFGSDKD PFPKGVIPLT AIEMTRSSKD NKFQVITGQR
360 370 380 390 400
VFVFRTESEA QRDLWCSTLQ SCLKEQRLLG HPRPPHPPRP LRTGTLELRG
410 420 430 440 450
HKAKVFAALI PGELALYKSE QAFSLGIGIC FIELQGCSVR ETKSRSFDLL
460 470 480 490 500
TPHRCFSFTA ESGGARQSWA AALQEAVTET LSDYEVAEKV WSNPANRHCA
510 520 530 540 550
DCRASRPDWA AVNLGVVICK QCAGQHRALG SGISKVQSLK LDTSVWSNEI
560 570 580 590 600
VQLFIVLGND RANCFWAGAL PPGEGLHPDS APGPRGEFIS RKYKLGLFRK
610 620 630 640 650
PHPRHPDHSQ LLQALCAAMA GPNLLKNMAQ LLCVETSEGE EPLSPSALNG
660 670 680 690 700
SLLSLLPSDS PGVYNEVVVP ATYRGFLYCG SISNKAGAPP LRRGRDAPPR
710 720 730 740 750
LWCVLGAALE MFASESSPEP LSLLQPQDIV CLGVSPPPAD PGDLDRFPFS
760 770 780 790 800
FELILTGGRI QHFATDGADS LEAWISAVGK WFSPLSCHQL LGPGLLRMGR
810 820 830 840 850
LWLRSPSHAG LAPGLWLSGF GLLRGDHLFL CPAPGPGPPA PEDMVHLRRL
860 870 880 890 900
QEISVVSAAD TPDKKEHLVL VETGRTLYLQ GEGRLDFAAW NTAIGGAAGG
910 920 930 940 950
GGTGLQEQQM SRGDIPIIVD ACISFVTQHG LRLEGVYRKG GARARSLRLL
960 970 980 990 1000
AEFRRDARSV KLRPREHFVE DVTDTLKRFF RELDDPVTSA RLLPRWREAA
1010 1020 1030 1040 1050
ELSQKNQRLE KYKEVISCLP RVNRRTLATL IGHLYRVQKC ASLNQMCTRN
1060 1070 1080 1090 1100
LALLFAPSVF QTDGRGEHEV RVLQELIDGY ISVFDIDSDQ AAQIDLEVSL
1110 1120 1130 1140 1150
ITTWKDVQLS QAGDLIMEVY IEQQLPDNCV TLKVSPTLTA EELTNQVLEM
1160 1170 1180 1190 1200
RGAASGTDLW VTFEILEHGE LERPLHPKEK VLEQALQWCQ LPEPCSASLL
1210 1220 1230 1240 1250
LRKVSMAHAG CLFTGVRRES PRVGLLRCRE EPPRLLGNRF QERFFLVRGR
1260 1270 1280 1290 1300
CLLLLKEKKS SKPEREWSLE GAKVYLGIRK KLKPPTLWGF TLILEKMHLC
1310 1320 1330 1340 1350
LSCMDEEEMW DWTTSILKAQ HDDQQSVVLR RRSSSDLARQ KFGTMPLLPI
1360 1370 1380 1390 1400
RGDDSGATLL SANQTLRRLH NRRTLSMFFP MKSPQGSVEE QDELEEPVYE
1410 1420 1430 1440 1450
EPVYEEVGAF PELTKDTTFS STWEWSAKSD PSLTSQRSFD QPPLSKASML
1460 1470 1480 1490 1500
GHEERIPDPP PGPPSKSSSQ ARGSLEEQLL QELNNLILRK GEPASCPESS
1510 1520 1530
SQPTSPQAPS PTSLPTPTPS LPTQPPCTSN PPSSQPLT
Length:1,538
Mass (Da):169,741
Last modified:July 27, 2011 - v3
Checksum:iD7E55C57FD4B83ED
GO
Isoform 2 (identifier: Q8R5G7-2) [UniParc]FASTAAdd to basket

Also known as: ARAP3 delta-SAM

The sequence of this isoform differs from the canonical sequence as follows:
     1-78: Missing.

Show »
Length:1,460
Mass (Da):161,171
Checksum:iC00130DB36CA8116
GO
Isoform 3 (identifier: Q8R5G7-3) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     1-618: Missing.
     1446-1538: KASMLGHEER...SNPPSSQPLT → GPSPFVCTMKSLKEGEG

Note: No experimental confirmation available.
Show »
Length:844
Mass (Da):94,043
Checksum:i5742FA191BD0F796
GO

Sequence cautioni

The sequence AAL78678.1 differs from that shown. Reason: Erroneous initiation. Curated
The sequence BAD90507.1 differs from that shown. Reason: Erroneous initiation. Curated

Experimental Info

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Sequence conflicti144 – 1441Q → H in AAH68145 (PubMed:15489334).Curated
Sequence conflicti167 – 1671R → Q in AAL78677 (PubMed:15546919).Curated
Sequence conflicti167 – 1671R → Q in AAL78678 (PubMed:15546919).Curated
Sequence conflicti1304 – 13041M → T in AAL78677 (PubMed:15546919).Curated
Sequence conflicti1304 – 13041M → T in AAL78678 (PubMed:15546919).Curated
Sequence conflicti1500 – 15001S → P in AAL78677 (PubMed:15546919).Curated
Sequence conflicti1500 – 15001S → P in AAL78678 (PubMed:15546919).Curated

Alternative sequence

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Alternative sequencei1 – 618618Missing in isoform 3. 1 PublicationVSP_015002Add
BLAST
Alternative sequencei1 – 7878Missing in isoform 2. 1 PublicationVSP_015003Add
BLAST
Alternative sequencei1446 – 153893KASML…SQPLT → GPSPFVCTMKSLKEGEG in isoform 3. 1 PublicationVSP_015004Add
BLAST

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AF469621 mRNA. Translation: AAL78677.1.
AF469622 mRNA. Translation: AAL78678.1. Different initiation.
AC121886 Genomic DNA. No translation available.
AC129315 Genomic DNA. No translation available.
BC068145 mRNA. Translation: AAH68145.1.
AK220486 mRNA. Translation: BAD90507.1. Different initiation.
CCDSiCCDS29196.1. [Q8R5G7-1]
RefSeqiNP_001192265.1. NM_001205336.1. [Q8R5G7-2]
NP_631945.2. NM_139206.2. [Q8R5G7-1]
XP_006525552.1. XM_006525489.1. [Q8R5G7-1]
UniGeneiMm.247642.

Genome annotation databases

EnsembliENSMUST00000042944; ENSMUSP00000035662; ENSMUSG00000024451. [Q8R5G7-1]
GeneIDi106952.
KEGGimmu:106952.
UCSCiuc008erq.2. mouse. [Q8R5G7-1]

Keywords - Coding sequence diversityi

Alternative splicing

Cross-referencesi

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AF469621 mRNA. Translation: AAL78677.1.
AF469622 mRNA. Translation: AAL78678.1. Different initiation.
AC121886 Genomic DNA. No translation available.
AC129315 Genomic DNA. No translation available.
BC068145 mRNA. Translation: AAH68145.1.
AK220486 mRNA. Translation: BAD90507.1. Different initiation.
CCDSiCCDS29196.1. [Q8R5G7-1]
RefSeqiNP_001192265.1. NM_001205336.1. [Q8R5G7-2]
NP_631945.2. NM_139206.2. [Q8R5G7-1]
XP_006525552.1. XM_006525489.1. [Q8R5G7-1]
UniGeneiMm.247642.

3D structure databases

ProteinModelPortaliQ8R5G7.
SMRiQ8R5G7. Positions 1-80.
ModBaseiSearch...
MobiDBiSearch...

Protein-protein interaction databases

IntActiQ8R5G7. 3 interactions.
STRINGi10090.ENSMUSP00000035662.

PTM databases

iPTMnetiQ8R5G7.
PhosphoSiteiQ8R5G7.

Proteomic databases

MaxQBiQ8R5G7.
PaxDbiQ8R5G7.
PRIDEiQ8R5G7.

Protocols and materials databases

Structural Biology KnowledgebaseSearch...

Genome annotation databases

EnsembliENSMUST00000042944; ENSMUSP00000035662; ENSMUSG00000024451. [Q8R5G7-1]
GeneIDi106952.
KEGGimmu:106952.
UCSCiuc008erq.2. mouse. [Q8R5G7-1]

Organism-specific databases

CTDi64411.
MGIiMGI:2147274. Arap3.
RougeiSearch...

Phylogenomic databases

eggNOGiKOG1117. Eukaryota.
COG5347. LUCA.
GeneTreeiENSGT00760000118874.
HOGENOMiHOG000246988.
HOVERGENiHBG069114.
InParanoidiQ8R5G7.
KOiK12490.
OMAiHEGIYRK.
OrthoDBiEOG7F24T4.
TreeFamiTF105769.

Enzyme and pathway databases

ReactomeiR-MMU-194840. Rho GTPase cycle.

Miscellaneous databases

NextBioi358482.
PROiQ8R5G7.
SOURCEiSearch...

Gene expression databases

BgeeiQ8R5G7.
GenevisibleiQ8R5G7. MM.

Family and domain databases

Gene3Di1.10.150.50. 1 hit.
1.10.555.10. 1 hit.
2.30.29.30. 3 hits.
InterProiIPR001164. ArfGAP.
IPR011993. PH_dom-like.
IPR001849. PH_domain.
IPR000159. RA_dom.
IPR008936. Rho_GTPase_activation_prot.
IPR000198. RhoGAP_dom.
IPR001660. SAM.
IPR013761. SAM/pointed.
IPR029071. Ubiquitin-rel_dom.
[Graphical view]
PfamiPF01412. ArfGap. 1 hit.
PF00169. PH. 2 hits.
PF00788. RA. 1 hit.
PF00620. RhoGAP. 1 hit.
PF07647. SAM_2. 1 hit.
[Graphical view]
PRINTSiPR00405. REVINTRACTNG.
SMARTiSM00105. ArfGap. 1 hit.
SM00233. PH. 5 hits.
SM00324. RhoGAP. 1 hit.
SM00454. SAM. 1 hit.
[Graphical view]
SUPFAMiSSF47769. SSF47769. 1 hit.
SSF48350. SSF48350. 1 hit.
SSF50729. SSF50729. 5 hits.
SSF54236. SSF54236. 1 hit.
PROSITEiPS50115. ARFGAP. 1 hit.
PS50003. PH_DOMAIN. 3 hits.
PS50200. RA. 1 hit.
PS50238. RHOGAP. 1 hit.
PS50105. SAM_DOMAIN. 1 hit.
[Graphical view]
ProtoNetiSearch...

Publicationsi

« Hide 'large scale' publications
  1. "ARAP3 is transiently tyrosine phosphorylated in cells attaching to fibronectin and inhibits cell spreading in a RhoGAP-dependent manner."
    Stacey T.T.I., Nie Z., Stewart A., Najdovska M., Hall N.E., He H., Randazzo P.A., Lock P.
    J. Cell Sci. 117:6071-6084(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORMS 1 AND 2), FUNCTION, MUTAGENESIS OF ARG-938; TYR-1399 AND TYR-1404, PHOSPHORYLATION AT TYR-1399 AND TYR-1404, SUBCELLULAR LOCATION.
    Strain: NIH Swiss.
    Tissue: Embryo.
  2. Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
    Strain: C57BL/6J.
  3. "The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC)."
    The MGC Project Team
    Genome Res. 14:2121-2127(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 1).
    Strain: C57BL/6J.
    Tissue: Brain.
  4. "Prediction of the coding sequences of mouse homologues of KIAA gene. The complete nucleotide sequences of mouse KIAA-homologous cDNAs identified by screening of terminal sequences of cDNA clones randomly sampled from size-fractionated libraries."
    Okazaki N., Kikuno R.F., Ohara R., Inamoto S., Nagase T., Ohara O., Koga H.
    Submitted (FEB-2005) to the EMBL/GenBank/DDBJ databases
    Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 619-1538 (ISOFORM 3).
    Tissue: Brain.
  5. "Quantitative time-resolved phosphoproteomic analysis of mast cell signaling."
    Cao L., Yu K., Banh C., Nguyen V., Ritz A., Raphael B.J., Kawakami Y., Kawakami T., Salomon A.R.
    J. Immunol. 179:5864-5876(2007) [PubMed] [Europe PMC] [Abstract]
    Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT TYR-1404, IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS].
    Tissue: Mast cell.
  6. Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT THR-1344, IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS].
    Tissue: Brown adipose tissue, Heart, Kidney, Lung and Spleen.

Entry informationi

Entry nameiARAP3_MOUSE
AccessioniPrimary (citable) accession number: Q8R5G7
Secondary accession number(s): E9QMK7
, Q5DTN4, Q6NVF1, Q8R5G6
Entry historyi
Integrated into UniProtKB/Swiss-Prot: August 16, 2005
Last sequence update: July 27, 2011
Last modified: April 13, 2016
This is version 131 of the entry and version 3 of the sequence. [Complete history]
Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programChordata Protein Annotation Program

Miscellaneousi

Keywords - Technical termi

Complete proteome, Reference proteome

Documents

  1. MGD cross-references
    Mouse Genome Database (MGD) cross-references in UniProtKB/Swiss-Prot
  2. SIMILARITY comments
    Index of protein domains and families

Similar proteinsi

Links to similar proteins from the UniProt Reference Clusters (UniRef) at 100%, 90% and 50% sequence identity:
100%UniRef100 combines identical sequences and sub-fragments with 11 or more residues from any organism into one UniRef entry.
90%UniRef90 is built by clustering UniRef100 sequences that have at least 90% sequence identity to, and 80% overlap with, the longest sequence (a.k.a seed sequence).
50%UniRef50 is built by clustering UniRef90 seed sequences that have at least 50% sequence identity to, and 80% overlap with, the longest sequence in the cluster.