CDH13 cadherin 13 [Homo sapiens (human)] - Gene

Summary

Official Symbol: CDH13provided by HGNC
Official Full Name: cadherin 13provided by HGNC
Primary source: HGNC:HGNC:1753
See related: Ensembl:ENSG00000140945 MIM:601364; AllianceGenome:HGNC:1753
Gene type: protein coding
RefSeq status: REVIEWED
Organism: Homo sapiens
Lineage: Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia; Eutheria; Euarchontoglires; Primates; Haplorrhini; Catarrhini; Hominidae; Homo
Also known as: CDHH; P105
Summary: This gene encodes a member of the cadherin superfamily. The encoded protein is localized to the surface of the cell membrane and is anchored by a GPI moiety, rather than by a transmembrane domain. The protein lacks the cytoplasmic domain characteristic of other cadherins, and so is not thought to be a cell-cell adhesion glycoprotein. This protein acts as a negative regulator of axon growth during neural differentiation. It also protects vascular endothelial cells from apoptosis due to oxidative stress, and is associated with resistance to atherosclerosis. The gene is hypermethylated in many types of cancer. Alternative splicing results in multiple transcript variants encoding different isoforms. [provided by RefSeq, May 2011]
Expression: Broad expression in endometrium (RPKM 22.0), heart (RPKM 17.7) and 16 other tissues See more
Orthologs: mouse all
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Genomic context

Location:: 16q23.3

Exon count:: 20

Annotation release	Status	Assembly	Chr	Location
RS_2024_08	current	GRCh38.p14 (GCF_000001405.40)	16	NC_000016.10 (82626969..83800640)
RS_2024_08	current	T2T-CHM13v2.0 (GCF_009914755.1)	16	NC_060940.1 (88691276..89866370)
RS_2024_09	previous assembly	GRCh37.p13 (GCF_000001405.25)	16	NC_000016.9 (82660574..83834245)

Chromosome 16 - NC_000016.10 Genomic Context describing neighboring genes

Genomic regions, transcripts, and products

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Go to reference sequence details

Genomic Sequence:

Go to nucleotide: Graphics FASTA GenBank

Expression

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See details

Project title: HPA RNA-seq normal tissues HPA RNA-seq normal tissues
Description: RNA-seq was performed of tissue samples from 95 human individuals representing 27 different tissues in order to determine tissue-specificity of all protein-coding genes
BioProject: PRJEB4337
Publication: PMID 24309898
Analysis date: Wed Apr 4 07:08:55 2018

Bibliography

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GeneRIFs: Gene References Into Functions

What's a GeneRIF?

Hsa_circ_0000119 promoted ovarian cancer development via enhancing the methylation of CDH13 by sponging miR-142-5p.
Title: Hsa_circ_0000119 promoted ovarian cancer development via enhancing the methylation of CDH13 by sponging miR-142-5p.
Cadherin-13 is a critical regulator of GABAergic modulation in human stem-cell-derived neuronal networks.
Title: Cadherin-13 is a critical regulator of GABAergic modulation in human stem-cell-derived neuronal networks.
A Common CDH13 Variant Is Associated with Low Agreeableness and Neural Responses to Working Memory Tasks in ADHD.
Title: A Common CDH13 Variant Is Associated with Low Agreeableness and Neural Responses to Working Memory Tasks in ADHD.
Genome-Wide Association Study on Adiponectin-Mediated Suppression of HDL-C Levels in Taiwanese Individuals Identifies Functional Haplotypes in CDH13.
Title: Genome-Wide Association Study on Adiponectin-Mediated Suppression of HDL-C Levels in Taiwanese Individuals Identifies Functional Haplotypes in CDH13.
Interactive association between dietary fat and sex on CDH13 cg02263260 methylation.
Title: Interactive association between dietary fat and sex on CDH13 cg02263260 methylation.
Revisiting the multiple roles of T-cadherin in health and disease.
Title: Revisiting the multiple roles of T-cadherin in health and disease.
Association of CDH13 Gene Polymorphism and Metabolic Syndrome in Gambian Population.
Title: Association of CDH13 Gene Polymorphism and Metabolic Syndrome in Gambian Population.
microRNA-377-3p downregulates the oncosuppressor T-cadherin in colorectal adenocarcinoma cells.
Title: microRNA-377-3p downregulates the oncosuppressor T-cadherin in colorectal adenocarcinoma cells.
The immunohistochemical expression of von Willebrand factor, T-cadherin, and Caveolin-1 is increased in kidney allograft biopsies with antibody-mediated injury.
Title: The immunohistochemical expression of von Willebrand factor, T-cadherin, and Caveolin-1 is increased in kidney allograft biopsies with antibody-mediated injury.
CDH13 and LPHN3 Gene Polymorphisms in Attention-Deficit/Hyperactivity Disorder: Their Relation to Clinical Characteristics.
Title: CDH13 and LPHN3 Gene Polymorphisms in Attention-Deficit/Hyperactivity Disorder: Their Relation to Clinical Characteristics.

Submit: New GeneRIF Correction See all GeneRIFs (151)

Phenotypes

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Find tests for this gene in the NIH Genetic Testing Registry (GTR)

Review eQTL and phenotype association data in this region using PheGenI

EBI GWAS Catalog

Description
A genome-wide association study in 19 633 Japanese subjects identified LHX3-QSOX2 and IGF1 as adult height loci. EBI GWAS Catalog EBI GWAS CatalogPubMed
A genome-wide association study of chemotherapy-induced alopecia in breast cancer patients. EBI GWAS Catalog EBI GWAS CatalogPubMed
A genome-wide association study of total serum and mite-specific IgEs in asthma patients. EBI GWAS Catalog EBI GWAS CatalogPubMed
A genome-wide association study reveals a quantitative trait locus of adiponectin on CDH13 that predicts cardiometabolic outcomes. EBI GWAS Catalog EBI GWAS CatalogPubMed
A meta-analysis of genome-wide association studies for adiponectin levels in East Asians identifies a novel locus near WDR11-FGFR2. EBI GWAS Catalog EBI GWAS CatalogPubMed
Adiponectin concentrations: a genome-wide association study. EBI GWAS Catalog EBI GWAS CatalogPubMed
CDH13 gene coding T-cadherin influences variations in plasma adiponectin levels in the Japanese population. EBI GWAS Catalog EBI GWAS CatalogPubMed
Common genetic variation and performance on standardized cognitive tests. EBI GWAS Catalog EBI GWAS CatalogPubMed
Framingham Heart Study 100K Project: genome-wide associations for blood pressure and arterial stiffness. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genetic predictors of fibrin D-dimer levels in healthy adults. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide association analyses identifies a susceptibility locus for tuberculosis on chromosome 18q11.2. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide association scan of trait depression. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide association study for adiponectin levels in Filipino women identifies CDH13 and a novel uncommon haplotype at KNG1-ADIPOQ. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide association study identifies three novel genetic markers associated with elite endurance performance. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide association study of 14,000 cases of seven common diseases and 3,000 shared controls. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide association study of chronic periodontitis in a general German population. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide association study of d-amphetamine response in healthy volunteers identifies putative associations, including cadherin 13 (CDH13). EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide association study of smoking behaviours among Bangladeshi adults. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide association study on dimethylarginines reveals novel AGXT2 variants associated with heart rate variability but not with overall mortality. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide contribution of genotype by environment interaction to variation of diabetes-related traits. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide meta-analysis identifies new susceptibility loci for migraine. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide scan identifies CDH13 as a novel susceptibility locus contributing to blood pressure determination in two European populations. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genome-wide study of association and interaction with maternal cytomegalovirus infection suggests new schizophrenia loci. EBI GWAS Catalog EBI GWAS CatalogPubMed
Genomewide pharmacogenomic study of metabolic side effects to antipsychotic drugs. EBI GWAS Catalog EBI GWAS CatalogPubMed
Imputation-based meta-analysis of severe malaria in three African populations. EBI GWAS Catalog EBI GWAS CatalogPubMed
Loci associated with N-glycosylation of human immunoglobulin G show pleiotropy with autoimmune diseases and haematological cancers. EBI GWAS Catalog EBI GWAS CatalogPubMed
Molecular genetics of adult ADHD: converging evidence from genome-wide association and extended pedigree linkage studies. EBI GWAS Catalog EBI GWAS CatalogPubMed
Novel loci for adiponectin levels and their influence on type 2 diabetes and metabolic traits: a multi-ethnic meta-analysis of 45,891 individuals. EBI GWAS Catalog EBI GWAS CatalogPubMed
Preliminary evidence of genetic determinants of adiponectin response to fenofibrate in the Genetics of Lipid Lowering Drugs and Diet Network. EBI GWAS Catalog EBI GWAS CatalogPubMed
The CCND1 c.870G&gt;A polymorphism is a risk factor for t(11;14)(q13;q32) multiple myeloma. EBI GWAS Catalog EBI GWAS CatalogPubMed

Variation

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See variants in ClinVar

See studies and variants in dbVar

See Variation Viewer (GRCh37.p13)

See Variation Viewer (GRCh38)

Interactions

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Products	Interactant	Other Gene	Complex	Source	Pubs	Description

General gene information

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Markers

SHGC-78995 (e-PCR)
- UniSTS:101379

SHGC-30008 (e-PCR)
- UniSTS:1031

D16S2789 (e-PCR)
- UniSTS:153359

RH65167 (e-PCR)
- UniSTS:90393

GDB:568474 (e-PCR)
- UniSTS:157721

D16S3091 (e-PCR), detects polymorphism
- UniSTS:53785

D16S422 (e-PCR)
- UniSTS:27172

D16S2949 (e-PCR)
- UniSTS:17710

D16S2820 (e-PCR)
- UniSTS:153390

SHGC-53490 (e-PCR)
- UniSTS:57135

DXS7657 (e-PCR)
- UniSTS:45877

SHGC-61168 (e-PCR)
- UniSTS:78144

RH103793 (e-PCR)
- UniSTS:98118

SHGC-154183 (e-PCR)
- UniSTS:178003

SHGC-151059 (e-PCR)
- UniSTS:173145

D16S422 (e-PCR)
- UniSTS:35433

G17933 (e-PCR)
- UniSTS:75636

RH122324 (e-PCR)
- UniSTS:137846

D16S735 (e-PCR)
- UniSTS:149879

SHGC-105014 (e-PCR)
- UniSTS:169086

SHGC-37041 (e-PCR)
- UniSTS:33680

D6S961 (e-PCR)
- UniSTS:18837

SHGC-142813 (e-PCR)
- UniSTS:171674

RH120935 (e-PCR)
- UniSTS:133794

SHGC-78198 (e-PCR)
- UniSTS:102054

D16S2641 (e-PCR)
- UniSTS:149422

D16S2692 (e-PCR)
- UniSTS:153262

RH92366 (e-PCR)
- UniSTS:85185

SHGC-149995 (e-PCR)
- UniSTS:177357

CDH13_7 (e-PCR)
- UniSTS:276174

RH121582 (e-PCR)
- UniSTS:134354

SHGC-152003 (e-PCR)
- UniSTS:177363

SHGC-156033 (e-PCR)
- UniSTS:183002

SHGC-60721 (e-PCR)
- UniSTS:41956

AFMB338ZB1 (e-PCR)
- UniSTS:14316

D16S402 (e-PCR)
- UniSTS:83180

D16S2756 (e-PCR)
- UniSTS:153326

SHGC-147944 (e-PCR)
- UniSTS:176114

SHGC-77957 (e-PCR)
- UniSTS:101364

Gene Ontology Provided by GOA

Function	Evidence Code	Pubs
enables adiponectin binding	ISS Inferred from Sequence or Structural Similarity more info
enables beta-catenin binding	IBA Inferred from Biological aspect of Ancestor more info
enables cadherin binding	IBA Inferred from Biological aspect of Ancestor more info
enables cadherin binding	IDA Inferred from Direct Assay more info	PubMed
enables calcium ion binding	IEA Inferred from Electronic Annotation more info
enables identical protein binding	IEA Inferred from Electronic Annotation more info
enables lipoprotein particle binding	IDA Inferred from Direct Assay more info	PubMed
enables low-density lipoprotein particle binding	IDA Inferred from Direct Assay more info	PubMed
enables protein binding	IPI Inferred from Physical Interaction more info	PubMed

Process	Evidence Code	Pubs
involved_in Rac protein signal transduction	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in Rho protein signal transduction	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in adherens junction organization	IBA Inferred from Biological aspect of Ancestor more info
involved_in calcium-dependent cell-cell adhesion via plasma membrane cell adhesion molecules	IBA Inferred from Biological aspect of Ancestor more info
involved_in calcium-dependent cell-cell adhesion via plasma membrane cell adhesion molecules	IDA Inferred from Direct Assay more info	PubMed
involved_in cell migration	IBA Inferred from Biological aspect of Ancestor more info
involved_in cell morphogenesis	IBA Inferred from Biological aspect of Ancestor more info
involved_in cell-cell adhesion mediated by cadherin	IBA Inferred from Biological aspect of Ancestor more info
involved_in cell-cell junction assembly	IBA Inferred from Biological aspect of Ancestor more info
involved_in endothelial cell migration	IDA Inferred from Direct Assay more info	PubMed
involved_in homophilic cell adhesion via plasma membrane adhesion molecules	IDA Inferred from Direct Assay more info	PubMed
involved_in keratinocyte proliferation	IDA Inferred from Direct Assay more info	PubMed
involved_in lamellipodium assembly	IDA Inferred from Direct Assay more info	PubMed
involved_in localization within membrane	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in low-density lipoprotein particle mediated signaling	IDA Inferred from Direct Assay more info	PubMed
involved_in negative regulation of cell adhesion	IDA Inferred from Direct Assay more info	PubMed
involved_in negative regulation of cell population proliferation	IDA Inferred from Direct Assay more info	PubMed
involved_in positive regulation of calcium-mediated signaling	IDA Inferred from Direct Assay more info	PubMed
involved_in positive regulation of cell migration	IDA Inferred from Direct Assay more info	PubMed
involved_in positive regulation of cell-matrix adhesion	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in positive regulation of endothelial cell proliferation	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in positive regulation of positive chemotaxis	IDA Inferred from Direct Assay more info	PubMed
involved_in positive regulation of smooth muscle cell proliferation	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in positive regulation of transcription by RNA polymerase II	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in regulation of endocytosis	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in regulation of epidermal growth factor receptor signaling pathway	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in sprouting angiogenesis	IDA Inferred from Direct Assay more info	PubMed

Component	Evidence Code	Pubs
located_in GABA-ergic synapse	IEA Inferred from Electronic Annotation more info
is_active_in adherens junction	IBA Inferred from Biological aspect of Ancestor more info
part_of catenin complex	IBA Inferred from Biological aspect of Ancestor more info
located_in caveola	IDA Inferred from Direct Assay more info	PubMed
located_in collagen-containing extracellular matrix	HDA more info	PubMed
located_in cytoplasm	IDA Inferred from Direct Assay more info	PubMed
located_in external side of plasma membrane	IEA Inferred from Electronic Annotation more info
located_in extracellular exosome	HDA more info	PubMed
located_in extracellular region	HDA more info	PubMed
located_in extracellular space	IDA Inferred from Direct Assay more info	PubMed
located_in focal adhesion	HDA more info	PubMed
located_in neuron projection	IDA Inferred from Direct Assay more info	PubMed
located_in perinuclear region of cytoplasm	IEA Inferred from Electronic Annotation more info
located_in plasma membrane	IDA Inferred from Direct Assay more info	PubMed
located_in plasma membrane	TAS Traceable Author Statement more info

General protein information

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Preferred Names: cadherin-13

Names: H-cadherin (heart); T-cad; T-cadherin; cadherin 13, H-cadherin (heart); heart cadherin

NP_001207417.1

EC 3.6.1.3

NP_001207418.1

EC 3.6.1.3

NP_001207419.1

EC 3.6.1.3

NP_001207420.1

EC 3.6.1.3

NP_001207421.1

EC 3.6.1.3

NP_001248.1

EC 3.6.1.3

XP_011521106.1

EC 3.6.1.3

XP_016878337.1

EC 3.6.1.3

XP_016878338.1

EC 3.6.1.3

XP_054235331.1

EC 3.6.1.3

XP_054235332.1

EC 3.6.1.3

XP_054235333.1

EC 3.6.1.3

NCBI Reference Sequences (RefSeq)

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RefSeqs maintained independently of Annotated Genomes

These reference sequences exist independently of genome builds. Explain

These reference sequences are curated independently of the genome annotation cycle, so their versions may not match the RefSeq versions in the current genome build. Identify version mismatches by comparing the version of the RefSeq in this section to the one reported in Genomic regions, transcripts, and products above.

Genomic

NG_052819.2 RefSeqGene

Range

5002..1178673

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GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

NM_001220488.2 → NP_001207417.1 cadherin-13 isoform 2

Status: REVIEWED

Description

Transcript Variant: This variant (2) includes an alternate exon in the 5' coding region, compared to variant 1. It encodes isoform 2, which is longer and has a distinct N-terminus, compared to isoform 1.

Source sequence(s)

AC009119, AC092351, AK304733, BG911233, BU627784, CB321814, CF618908, CV024366, DA561809, DA685616, DB003683, DC376661, R41787

Consensus CDS

CCDS58485.1

UniProtKB/TrEMBL

B7Z9B1

Related

ENSP00000268613.10, ENST00000268613.14

Conserved Domains (5) summary

cd11304
Location:535 → 628

Cadherin_repeat; Cadherin tandem repeat domain

smart00112
Location:208 → 290

CA; Cadherin repeats

pfam00028
Location:415 → 516

Cadherin; Cadherin domain

pfam08758
Location:74 → 159

Cadherin_pro; Cadherin prodomain like

cl09101
Location:636 → 727

E_set; Early set domain associated with the catalytic domain of sugar utilizing enzymes at either the N or C terminus
NM_001220489.2 → NP_001207418.1 cadherin-13 isoform 3 precursor

See identical proteins and their annotated locations for NP_001207418.1

Status: REVIEWED

Description

Transcript Variant: This variant (3) lacks an in-frame exon in the coding region, compared to variant 1. It encodes isoform 3, which is shorter than isoform 1.

Source sequence(s)

AC009119, AK298612, BG911233, BU627784, CB321814, CF618908, DA685616, DC376661, R41787

Consensus CDS

CCDS58487.1

UniProtKB/TrEMBL

B7Z9B1

Related

ENSP00000394557.3, ENST00000428848.7

Conserved Domains (5) summary

cd11304
Location:449 → 542

Cadherin_repeat; Cadherin tandem repeat domain

smart00112
Location:122 → 204

CA; Cadherin repeats

pfam00028
Location:329 → 430

Cadherin; Cadherin domain

pfam08758
Location:27 → 112

Cadherin_pro; Cadherin prodomain like

cl09101
Location:550 → 641

E_set; Early set domain associated with the catalytic domain of sugar utilizing enzymes at either the N or C terminus
NM_001220490.2 → NP_001207419.1 cadherin-13 isoform 4

Status: REVIEWED

Description

Transcript Variant: This variant (4) lacks an exon in the coding region resulting in use of a downstream start codon, compared to variant 1. It encodes isoform 4, which is shorter and has a distinct N-terminus, compared to isoform 1.

Source sequence(s)

AC009119, AK295886, BU162665, BU627784, CB321814, CF618908, DA685616, DC376661, R41787

UniProtKB/TrEMBL

B7Z3H7

Conserved Domains (3) summary

cd11304
Location:234 → 327

Cadherin_repeat; Cadherin tandem repeat domain

pfam00028
Location:114 → 215

Cadherin; Cadherin domain

cl29226
Location:335 → 426

Cadherin; Cadherin domain
NM_001220491.2 → NP_001207420.1 cadherin-13 isoform 5 precursor

See identical proteins and their annotated locations for NP_001207420.1

Status: REVIEWED

Description

Transcript Variant: This variant (5) lacks several coding exons and uses an alternate 3' terminal exon, compared to variant 1. It encodes isoform 5, which is shorter and has a distinct C-terminus, compared to isoform 1.

Source sequence(s)

AC087189, DC376661, EU190358

Consensus CDS

CCDS56010.1

UniProtKB/Swiss-Prot

P55290

Related

ENSP00000408632.3, ENST00000431540.7

Conserved Domains (1) summary

pfam08758
Location:27 → 112

Cadherin_pro; Cadherin prodomain like
NM_001220492.2 → NP_001207421.1 cadherin-13 isoform 6 precursor

See identical proteins and their annotated locations for NP_001207421.1

Status: REVIEWED

Description

Transcript Variant: This variant (6) lacks several coding exons and includes two alternate exons at the 3' end, compared to variant 1. It encodes isoform 6, which is shorter and has a distinct C-terminus, compared to isoform 1.

Source sequence(s)

DC376661, EU190357

Consensus CDS

CCDS56009.1

UniProtKB/Swiss-Prot

P55290

Related

ENSP00000456491.1, ENST00000565636.5

Conserved Domains (1) summary

pfam08758
Location:27 → 112

Cadherin_pro; Cadherin prodomain like
NM_001257.5 → NP_001248.1 cadherin-13 isoform 1 preproprotein

See identical proteins and their annotated locations for NP_001248.1

Status: REVIEWED

Description

Transcript Variant: This variant (1) represents the predominant transcript and encodes isoform 1.

Source sequence(s)

AC009119, BC028624, BG911233, BI756586, BU627784, CA445317, CN263227, DC376661, R41787

Consensus CDS

CCDS58486.1

UniProtKB/Swiss-Prot

A8W476, A8W477, B7Z590, C9JRI6, J3KN62, P55290, Q6GTW4, Q8TBX3

UniProtKB/TrEMBL

B7Z9B1

Related

ENSP00000479395.1, ENST00000567109.6

Conserved Domains (5) summary

cd11304
Location:488 → 581

Cadherin_repeat; Cadherin tandem repeat domain

smart00112
Location:161 → 243

CA; Cadherin repeats

pfam00028
Location:368 → 469

Cadherin; Cadherin domain

pfam08758
Location:27 → 112

Cadherin_pro; Cadherin prodomain like

cl09101
Location:589 → 680

E_set; Early set domain associated with the catalytic domain of sugar utilizing enzymes at either the N or C terminus

RefSeqs of Annotated Genomes: GCF_000001405.40-RS_2024_08

The following sections contain reference sequences that belong to a specific genome build. Explain

Reference GRCh38.p14 Primary Assembly

Genomic

NC_000016.10 Reference GRCh38.p14 Primary Assembly

Range

82626969..83800640

Download

GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

XM_017022848.3 → XP_016878337.1 cadherin-13 isoform X2
XM_017022849.3 → XP_016878338.1 cadherin-13 isoform X3
XM_011522804.4 → XP_011521106.1 cadherin-13 isoform X1

Conserved Domains (4) summary

cd11304
Location:387 → 480

Cadherin_repeat; Cadherin tandem repeat domain

smart00112
Location:60 → 142

CA; Cadherin repeats

pfam00028
Location:267 → 368

Cadherin; Cadherin domain

cl09101
Location:488 → 579

E_set; Early set domain associated with the catalytic domain of sugar utilizing enzymes at either the N or C terminus