Carbohydrate-Binding Module Family 13

Known Activities	Modules of approx. 150 residues which always appear as a threefold internal repeat. The only apparent exception to this, xylanase II of Actinomadura sp. FC7 (GenBank U08894), is in fact not completely sequenced. These modules were first identified in several plant lectins such as ricin or agglutinin of Ricinus communis which bind galactose residues. The three-dimensional structure of a plant lectin has been determined and displays a pseudo-threefold symmetry in accord with the observed sequence threefold repeat. These modules have since been found in a number of other proteins of various functions including glycoside hydrolases and glycosyltransferases. While in the plant lectins this module binds mannose, binding to xylan has been demonstrated in the Streptomyces lividans xylanase A and arabinofuranosidase B. Binding to GalNAc has been shown for the corresponding module of GalNAc transferase 4. For the other proteins, the binding specificity of these modules has not been established. The pseudo three-fold symmetry of the CBM13 module has now been confirmed in the 3-D structure of the intact, two-domain, xylanase of Streptomyces olivaceoviridis.
3D Structure Status	b-trefoil
Note	Previously known as cellulose-binding domain family XIII (CBD XIII).
External resources	HOMSTRAD;
Statistics	GenBank accession (2377); Uniprot accession (891); PDB accession (85); 3D entries (41); cryst (1)

Summary

All (1793) Archaea (13) Bacteria (1361) Eukaryota (401) Viruses (10) unclassified (8) Structure (41 - 1 cryst) Characterized (87)


Bacteria
Protein Name	EC#	Organism	GenBank	Uniprot	PDB/3D
xylanase II (XylII) (fragment)	3.2.1.8	Actinomadura sp. FC7	AAA17888.1 AAO97631.1 AAQ80272.1	Q59139
β-agarase (AgaB34)	3.2.1.81	Agarivorans albus YKW-34	ABW77762.1	A8W969
β-agarase (AgaJA3;Hz4)	3.2.1.81	Agarivorans sp. JA-1 / HZ105	ADY17921.1
β-agarase (AgaA)	3.2.1.81	Agarivorans sp. LQ48	ACM50513.1	C9WIW7
β-1,3-glucanase (GlcI)	3.2.1.39	Arthrobacter sp. YCWD3	BAA04892.1	Q59146
β-1,3(4)-glucanase M (BglM)	3.2.1.39 3.2.1.6 3.2.1.73	Bacillus circulans IAM1165	BAC06195.1	Q8KKH3
pectate lyase	4.2.2.2	Bacillus sp. AAI12	AAE59747.1 AAS29294.1
feruloyl esterase I (FaeI;CGSCsYakCAS_18248) (Fae1A)	3.1.1.73	Cellulosilyticum ruminicola CGMCC 1.5065	ACZ98598.1	D2KFJ6
β-1,3-glucanase (BglI)	3.2.1.39	Cellulosimicrobium cellulans 73 14 / ATCC21606 / DSM 10297	AAA25520.1	P22222
β-1,3-glucanase II (BglII)	3.2.1.39	Cellulosimicrobium cellulans LL-G109 / DSM10297 / DK-1	AAC38290.1 AEK80404.1	O68641
exo-β-1,3-galactanase (Ct1,3Gal43A;Cthe_0661) (1,3Gal43A)	3.2.1.145	Clostridium thermocellum ATCC 27405	ABN51896.1	A3DD67	3VSF[A,B,C,D,E,F] 3VSZ[A,B,C,D,E,F] 3VT0[A,B,C,D,E,F] 3VT1[A,B,C,D,E,F] 3VT2[A,B,C,D,E,F]
xylanase (M6;CtM6;Cthe_2193) (arabinoxylan-specific) (Xyl5A)	3.2.1.-	Clostridium thermocellum ATCC 27405	ABN53395.1	A3DHG6
β-agarase (AgaY) (extracellular)	3.2.1.81	Janthinobacterium sp. SY12	ABO93616.2	A4UMV9
pectate lyase B (PelB)	4.2.2.2	Pseudoalteromonas haloplanktis 505	AAF86343.1 AAF86343.2	Q9KGY6
β-agarase (AgaA) (extracellular)	3.2.1.81	Pseudoalteromonas sp. CY24	AAN39119.2	Q8GB62
β-agarase (AgaA6)	3.2.1.81	Pseudoalteromonas sp. JT-6	ACF40223.1	C0J1J2
β-agarase (PagA)	3.2.1.81	Pseudomonas sp. BK1 SK38 / BK1
β-porphyranase (AagA)	3.2.1.178	Pseudomonas sp. ND137	BAB79291.1	Q8VUT1
alginate lyase / poly(β-mannuronate) lyase (Sde_2547) (Alg7D)	4.2.2.3	Saccharophagus degradans 2-40	ABD81807.1 ACK10570.1 ACK10582.1	Q21HM2
endo-β-1,3-xylanase (Xyl30)	3.2.1.32	Streptomyces avermitilis CECT3339	AAD32560.1	Q9X584
exo-β-1,3-galactanase (Sa1,3Gal43A;SAV2109) (1,3Gal43A)	3.2.1.145	Streptomyces avermitilis MA-4680	BAC69820.1 NP_823285.1
β-L-arabinopyranosidase (AgaA2;SaArap27A;SAV_2186) (Arap27A)	3.2.1.88	Streptomyces avermitilis MA-4680	BAC69897.1 NP_823362.1		3A21[A,B] 3A22[A,B] 3A23[A,B]
bifunctional xylanase / α-L-arabinofuranosidase (Xyl23)	3.2.1.55 3.2.1.8	Streptomyces chattanoogensis UAH23	AAD32559.2 ABC05324.1	Q9X583
xylanase A (XlnA;SCO5931;SC10A5.36c)	3.2.1.8	Streptomyces coelicolor A3(2)	CAA16468.1 CAD55241.1 NP_733679.1	Q8CJQ1
xylanase (Xyn10)	3.2.1.8	Streptomyces fradiae var. k11	ABR14270.1	A7TVD4
α-L-arabinofuranosidase B (AbfB)	3.2.1.55	Streptomyces lividans 1326	AAC26524.1	P96463
xylanase A (XlnA) (Xyn10A)	3.2.1.8	Streptomyces lividans 1326 / IAF18	AAC26525.1 AAO97633.1	P26514	1KNL[A] 1KNM[A] 1KNN[A] 1MC9[A]
β-1,4-xylanase (XynA;FXYN;SoXyn10A) (Xyn10A)	3.2.1.8	Streptomyces olivaceoviridis E-86		Q7SI98	1ISV[A,B] 1ISW[A,B] 1ISX[A,B] 1ISY[A,B] 1ISZ[A,B] 1IT0[A,B] 1V6U[A,B] 1V6V[A,B] 1V6W[A,B] 1V6X[A,B] 2D1Z[A,B] 2D20[A,B] 2D22[A,B] 2D23[A,B] 2D24[A,B] 1XYF[A]
exo-β-1,3-galactanase (SGalase2)	3.2.1.145	Streptomyces sp. 19(2012)	AFH55135.1
exo-β-1,3-galactanase (SGalase1) (partial)	3.2.1.145	Streptomyces sp. 19(2012)	AFH55134.1
endo-β-1,3-glucanase (BglS27)	3.2.1.39	Streptomyces sp. S27	ACO94508.1	C3VPJ1
xylanase (XynAS27)	3.2.1.8	Streptomyces sp. S27	ACF57946.1	C3RYK8
endo-β-1,4-xylanase (XynSW2;XynSW2A;XynSW2B)	3.2.1.8	Streptomyces sp. SWU10	BAK19338.1
xylanase A	3.2.1.8	Streptomyces thermocyaneoviolaceus	AAF04600.1	Q9RMM5
xylanase I (StXI)	3.2.1.8	Streptomyces thermoviolaceus OPC-520	BAA19777.1 BAD02382.1	Q76BV3
chitinase (Chi35)	3.2.1.14	Streptomyces thermoviolaceus OPC-520	BAA88833.1	Q9RHU5
endoxylanase (XynB) (Xyn10A)	3.2.1.8	Thermopolyspora flexuosa DSM43186 / ATCC35864	AAO97629.1 AAO97630.1 AAO97632.1 AAQ80270.1 AAQ80271.1 AAQ80273.1 AAS23966.1 AAS23967.1 AAS23969.1 CAD48748.1 CAD48748.2	Q8GMV6
β-agarase D (AgaD)	3.2.1.81	Vibrio sp. PO-303	BAF34350.1	Q05KJ8
β-agarase (AgaV)	3.2.1.81	Vibrio sp. V134	ABL06969.1	A1E2B6
Eukaryota
Protein Name	EC#	Organism	GenBank	Uniprot	PDB/3D
[Blood group A] α-N-acetylgalactosaminidase (NagA)	3.2.1.49	Acremonium sp. No.413	BAB08149.1	Q9HFB1
α-N-acetylgalactosaminidase / α-galactosidase (AglA;An06g00170)	3.2.1.22 3.2.1.49	Aspergillus niger CBS 513.88	CAK44933.1	A2QL72
α-galactosidase A (Ag1A;AglA)	3.2.1.22	Aspergillus niger N402	AAE69559.1 CAA44950.1	P28351
UDP-GalNAc: polypeptide N-acetylgalactosaminyltransferase 1 (GalNT1;ppGalNAcT1;GalNAc-T1)	2.4.1.41	Bos taurus	AAA30532.1 AAA68489.1 AAE24810.1 AAE24816.1 NP_803485.1	Q07537
ZK688.8 (gly3)	2.4.1.41	Caenorhabditis elegans Bristol N2	AAA28224.3 AAC13669.1 CCD62541.1 NP_498722.1	P34678
Y116F11B.12 (Gly-4)	2.4.1.41	Caenorhabditis elegans Bristol N2	AAC13670.1 CAC14394.1 CBA11615.1 NP_507850.1	Q8I136 Q8I136
H38K22.5 (Gly-6)	2.4.1.41	Caenorhabditis elegans Bristol N2	AAC13674.1 AAC13675.1 AAC13676.1 CAA19707.1 CAC42317.1 CAC42318.1 NP_001022644.1	O61394
Y46H3A.6 (gly-7)	2.4.1.41	Caenorhabditis elegans Bristol N2	AAC13677.1 AAF60620.1 CCD70652.1 NP_503512.1	O61397
GLY9 (Y47D3A.23)	2.4.1.41	Caenorhabditis elegans Bristol N2	AAC13679.1 CAB57897.1 CAI46621.1 NP_499451.1	Q9U2C4 Q9U2C4
UDP-GalNAc:polypeptide N-acetylgalactosaminyltransferase T3	2.4.1.41	Cryptosporidium parvum	AAQ95175.1 AAQ95175.2	Q6TBQ9
UDP-GalNAc:polypeptide N-acetylgalactosaminyltransferase T1	2.4.1.41	Cryptosporidium parvum	AAQ95174.1 AAQ95174.2	Q6TBR0
polypeptide N-acetylgalactosaminyltransferase T4 (CG31956;AT25481p1)	2.4.1.41	Drosophila melanogaster	AAN10370.1 AAN10370.2 AAN75751.1 AAQ56701.1 ACL82986.1 AER27632.1 NP_722910.1	B7Z006 Q8IA42 Q8IA42
polypeptide N-acetylgalactosaminyltransferase T3 (CG4445)	2.4.1.41	Drosophila melanogaster	AAD38630.1 AAF59298.1 NP_610256.1	Q9Y117
polypeptide N-acetylgalactosaminyltransferase T35A / l(2)35Aa (Pgant35A;CG7480)	2.4.1.41	Drosophila melanogaster	AAF44871.1 AAF53391.1 AAK66862.1 AAM62405.1 AAM62406.1 AEG66944.1 NP_652069.2	F5XVG5 Q8MVS5 Q8MVS5
polypeptide N-acetylgalactosaminyltransferase T7 (CG6394)	2.4.1.41	Drosophila melanogaster	AAF48851.1 AAM62412.1 AAN09470.1 AAQ56704.1 AAV37008.1 NP_573301.2 NP_728178.1	Q8MV48 Q8MV48
polypeptide N-acetylgalactosaminyltransferase T6 (CG2103)	2.4.1.41	Drosophila melanogaster	AAF47689.1 AAF47690.1 AAL29177.1 AAQ56703.1 AGB94037.1 NP_647749.2 NP_728779.1	Q6WV16
polypeptide N-acetylgalactosaminyltransferase T8 (CG7297)	2.4.1.41	Drosophila melanogaster	AAF49587.1 AAL48588.1 NP_648800.1	Q9VUT6
polypeptide N-acetylgalactosaminyltransferase T2 (CG3254)	2.4.1.41	Drosophila melanogaster	AAF51113.2 AAF51113.3 AAQ22499.1 AAQ56700.1 AAS64620.1 AAS64620.2 NP_608773.1	Q6WV19
polypeptide N-acetylgalactosaminyltransferase T5 (CG31651)	2.4.1.41	Drosophila melanogaster	AAF52218.2 AAL25377.1 AAQ56702.1 ABI31292.1 NP_608906.2	Q6WV17
polypeptide N-acetylgalactosaminyltransferase T1 (CG8182)	2.4.1.41	Drosophila melanogaster	AAF58130.1 AAG13184.1 AAM29416.1 AAM70974.1 AAQ56699.1 NP_611043.1 NP_725472.1	Q6WV20
UDP-GalNAc: polypeptide N-acetylgalactosaminyltransferase 2 (GALNTL2;GALNT7)	2.4.1.41	Homo sapiens	AAE72285.1 AAE73125.1 AAH14789.1 AAH30625.1 AAK63127.1 AAQ88808.1 ABA31473.1 ABA36473.1 ABA38055.1 ABA68732.1 ABA69735.1 ABA70117.1 ABA70240.1 ABC04061.1 ABC16096.1 ABE23551.1 ABE26114.1 ABE26387.1 ABH64624.1 ABH64747.1 ABH64904.1 ABH66133.1 ABH70952.1 ABH71075.1 ABH72589.1 ABH75704.1 ABH99489.1 ABI02594.1 ABI07395.1 ABI11043.1 ABI12263.1 ABJ19428.1 ABJ34423.1 ABL21883.1 ABX94001.1 ABX96294.1 ABX96604.1 ABY05068.1 ABY16007.1 ABY18804.1 ACC22268.1 ACC22839.1 ACC23718.1 ACC32509.1 ACC34084.1 ACE08795.1 ACG89706.1 ACK00002.1 ACM95112.1 ACN08875.1 ACP73867.1 ACS06493.1 ACS08649.1 ACW23860.1 BAD29961.1 BAG52928.1 CAD38585.1 CAD89983.1 CAG32963.1 CAK32357.1 NP_473451.2	F1LIP6 Q8N3T1
Polypeptide GalNAc transferase T13 (contains KIAA1918)	2.4.1.41	Homo sapiens	AAI01032.1 AAI01034.1 AAI01035.1 ACC22253.1 BAC54545.1 BAG54747.1 CAD44533.2 CAG32964.1 NP_443149.2	Q68VI8 Q8IUC8
UDP-GalNAc: polypeptide N-acetylgalactosaminyltransferase 14 (GalNT14)	2.4.1.41	Homo sapiens	AAH06269.1 AAH06269.2 AAH10659.1 AAQ89118.1 AAX88899.1 ABA66763.1 ABH82928.1 ABI02314.1 ABI12599.1 ABJ30825.1 ABJ38955.1 ABX91594.1 ABX91870.1 ABX92146.1 ABX92422.1 ABX92698.1 ABX93132.1 ABX93639.1 ABX94373.1 ABX95380.1 ABX95656.1 ABX95932.1 ABY04154.1 ABY04430.1 ABY04706.1 ABY05292.1 ABY16398.1 ABY17890.1 ABY18166.1 ABY18442.1 ACC25543.1 ACC26284.1 ACC36580.1 ACC36856.1 ACE07501.1 ACE09339.1 ACE10551.1 ACJ96114.1 ACN02380.1 ACN06790.1 ACP73505.1 ACP89033.1 ACR88983.1 ACR89535.1 ACW28625.1 BAB14227.1 BAB14795.1 BAC03634.1 BAC56889.1 BAG53701.1 BAH12861.1 CAA70505.3 CAA70505.4 CAD10224.1 CAD34770.1 CAD44543.1 CAE91230.1 CAE91506.1 NP_078848.2	Q96FL9
UDP-GalNAc:polypeptide N-acetylgalactosaminyltransferase (GalNAc-T11;ppGalNAcT11;GalNT11)	2.4.1.41	Homo sapiens	AAD45821.1 ACC22266.1 BAB15105.1 BAB15338.1 BAG54116.1 BAH12901.1 CAC79625.2 CAC79625.3 CAD38481.1 NP_071370.2	Q8NCW6 Q8NCW6
UDP-GalNAc:polypeptide α-N-acetylgalactosaminyltransferase T4 (GalNAc-T4;ppGalNAcT4;GALNT4)	2.4.1.41	Homo sapiens	AAH36390.1 AAN94203.1 AAN94205.1 BAG35722.1 BAG60038.1 CAA69875.1 NP_003765.1	Q8N4A0 Q8N4A0
UDP-GalNAc:polypeptide α-N-acetylgalactosaminyltransferase T1 (GalNAc-T1;ppGalNAcT1;GALNT1)	2.4.1.41	Homo sapiens	AAC50327.1 AAH38440.1 ACC22256.1 BAF85751.1 CAA59380.1 CAD44535.1 NP_065207.2	Q10472 Q10472
UDP-GalNAc:polypeptide α-N-acetylgalactosaminyltransferase T2 (GalNAc-T2;ppGalNAcT2;GalNT2)	2.4.1.41	Homo sapiens	AAH41120.1 ACC22257.1 ACS13744.1 BAF82737.1 BAH12448.1 BAH13288.1 CAA59381.1 CAC00585.2 CAI22902.1 CAI23447.1 NP_004472.1	Q10471	2FFU[A] 2FFV[A,B]
UDP-GalNAc:polypeptide α-N-acetylgalactosaminyltransferase T3 (ppGalNacT3;GALNAc-T3;GALNT3)	2.4.1.41	Homo sapiens	AAI13566.1 AAI13568.1 AAY14678.1 BAG36187.1 CAA63371.1 CAD44536.1 NP_004473.1	Q14435
UDP-GalNAc:polypeptide α-N-acetylgalactosaminyltransferase T6 (GalNAc-T6)	2.4.1.41	Homo sapiens	AAH35822.2 AAI14506.1 BAB15297.1 BAC11118.1 BAG52560.1 BAG59418.1 BAJ10977.1 CAA69876.1 CAF85889.1 CAM83953.1 CAM83954.1 NP_009141.1 NP_009141.2	D7UNW5 Q8NCL4 Q8NCL4
UDP-GalNAc:polypeptide N-acetylgalactosaminyltransferase 5 (ppGalNacT5;GALNT5)	2.4.1.41	Homo sapiens	AAF15313.1 AAI42677.1 AAP34404.1 ACC22260.1 BAF84843.1 CAB65104.1 CAD44537.1 NP_055383.1 XP_050509.1	A5PKZ1 Q7Z7M9 Q7Z7M9
UDP-GalNAc:polypeptide N-acetylgalactosaminyltransferase 7 (Galnt7)	2.4.1.41	Homo sapiens	AAH35303.1 AAH46129.1 AAH47468.1 BAG52155.1 CAB60270.1 CAC39783.1 CAD44538.1 CAI72154.1 NP_059119.1	B3KQU3 Q86SF2 Q86SF2
polypeptide GalNAc transferase 8 (GalNAc-T8)	2.4.1.41	Homo sapiens	CAD44539.1 NP_059113.1	Q9NY28
polypeptide N-acetylgalactosaminyltransferase 9 (GalNAc-T9)	2.4.1.41	Homo sapiens	AAH69354.1 AAH93817.1 AAH93817.2 AAH93819.1 AAH93819.2 AAM49722.1 AAT18857.1 ABE14200.1 BAB13699.2 BAG51429.1 BAG51570.1 CAD44540.1 NP_068580.1	Q9HCQ5 Q9HCQ5
UDP-GalNAc: polypeptide N-acetylgalactosaminyltransferase 10 (GalNT10;ppGalNAcT10;pp-GalNAc-T10)	2.4.1.41	Homo sapiens	AAH07224.2 AAH72450.1 AAH98444.1 AAP65372.1 AAP65373.1 AAP65378.1 AAP65379.1 ABA33148.1 ABA33149.1 ABA33154.1 ABA33155.1 BAB14676.1 BAC56890.1 BAG54441.1 CAB46378.1 CAD44532.1 CAE91109.1 NP_060010.2 NP_938080.1	Q86SR1 Q86SR1	2D7I[A] 2D7R[A]
UDP-GalNAc: polypeptide N-acetylgalactosaminyltransferase 12 (ppGalNAcT12;Galnt12)	2.4.1.41	Homo sapiens	AAH13945.1 ABA38425.1 BAB15027.1 BAC07181.1 BAD93346.1 CAC80100.2 CAD44541.1 CAG32962.1 CAI17042.1 NP_078918.3	Q8IXK2 Q8IXK2
UDP-GalNAc: polypeptide α-N-acetylgalactosaminyltransferase T1 (GalNAc-T1;ppGalNAcTase-1;Galnt1)	2.4.1.41	Mus musculus	AAB58477.1 AAH56215.1 AAH90962.1 BAE28617.1 NP_038842.2	O08912 O08912	1XHB[A]
Polypeptide GalNAc transferase T3 (GALNT3)	2.4.1.41	Mus musculus	AAB09579.1 AAH43331.1 BAE22551.1 CAM20333.1 CAM21878.1 NP_056551.1	P70419 P70419
Polypeptide GalNAc transferase T4	2.4.1.41	Mus musculus	AAB58301.1 AAH57882.1 ABA38430.1 BAC28317.1 BAE28303.1 BAE31844.1 NP_056552.1	O08832
UDP-GalNAc:polypeptide N-acetylgalactosaminyltransferase 7 (Galnt7)	2.4.1.41	Mus musculus	AAH07484.1 AAH49907.1 AAH52461.1 AAK37549.1 BAC28284.1 BAC29461.1 BAE30388.1 BAE30662.1 BAE31034.1 BAE31754.1 NP_653332.2	Q80VA0 Q80VA0
polypeptide N-acetylgalactosaminyltransferase T2 (Galnt2)	2.4.1.41	Mus musculus	AAH07172.1 AAH53063.1 AAH59818.1 AAK37548.1 BAC32790.1 BAE26246.1 BAE30483.1 NP_644678.1	Q6PB93
UDP-GalNAc:polypeptide N-acetylgalactosaminyltransferase GalNT11	2.4.1.41	Mus musculus	AAH11428.1 AAH21504.1 AAH34185.1 AAH36143.1 AAH36145.1 AAQ06668.1 BAC40096.1 BAE36442.1 CAC79626.1	Q921L8 Q921L8
Polypeptide GalNAc transferase T6	2.4.1.41	Mus musculus C57BL/6J	AAI19325.1 AAI19327.1 BAC25984.1 BAC33618.1 BAE32989.1 BAE35316.1 CAB55352.1 NP_766039.1	Q0VE84 Q8C7U7 Q8C7U7
Polypeptide GalNAc transferase T6	2.4.1.41	Rattus norvegicus	AAC99426.1 NP_075215.1	Q9R0C5
UDP-GalNAc: polypeptide N-acetylgalactosaminyltransferase T1 (GalNAc-T1;ppGalNAcT1)	2.4.1.41	Rattus norvegicus	AAC52511.1 NP_077349.1	Q10473
N-acetylgalactosaminyltransferase T5 (Galnt5)	2.4.1.41	Rattus norvegicus	AAC69708.1 NP_113984.1	O88422
UDP-GalNAc:polypeptide N-acetylgalactosaminyltransferase T9	2.4.1.41	Rattus norvegicus	AAK54498.1 EDM04502.1 NP_570098.1	Q925R7
polypeptide N-acetylgalactosaminyltransferase T1	2.4.1.41	Sus scrofa	BAA12800.1	Q29121
UDP-GalNAc:polypeptide N-acetylgalactosaminyltransferase T3 (ppGalNAc-T3)	2.4.1.41	Toxoplasma gondii	AAO20278.1 AAQ95173.1	Q6YBY0
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Footnotes

Experimental characterization exclusively covers enzyme activities at present. If you are aware of any missing experimental characterization that should be mentioned here, we encourage you to send your comment and support of the experimental evidence, including an electronic copy or appropriate link to the support material (i.e., article(s), thesis, database, etc), to cazy@afmb.univ-mrs.fr.