Pathway Human Homo sapiens

Asparagine N-linked glycosylation

R-HSA-446203 in Reactome release 97: under Post-translational protein modification, with 312 genes placed in it by the mapping files and 3 child pathways in the hierarchy.

The same number in the other species

Reactome writes the mouse and rat events it infers from a human pathway under the human number with the species token. A door opens only where that species' own list in this release holds the id: R-MMU-446203 (mouse), R-RNO-446203 (rat). Whether the event was inferred from this one is what the record says.

The record
The genes
Parents and children

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This page reads the pathway's record from its source while you read the rest; with scripts off the card stays at its loading line. Reload with scripts on to see it, or open the source's own page from the link in the card.

01The record

Reactome's own record of this pathway

What this tells you

The pathway list, the hierarchy and the genes on this page are read from the files Reactome publishes for Reactome release 97, built into this site on 2026-09-09: the pathway list, the hierarchy relationship file and the two gene mapping files. The record card is the one live read, Reactome's own record of this pathway.

Reactome's download page describes the mapping files: Mapping files link the source database identifier to the lowest level pathway diagram or subset of the pathway, all levels of the pathway hierarchy or database identifier to all reactions. [R13] The gene list here is the all-levels mapping of NCBI Gene ids, filled from the all-levels mapping of Ensembl ids where the NCBI file has no row for a gene; each row names which file it came from by the shape of its source id. Each row carries the evidence codes the file writes for it, as written and unranked, and both where the file writes both; no page of Reactome's documentation the survey read defines the codes, so this page does not expand them. A gene id the identity files do not name is kept as the source's own id with no door rather than turned into a symbol.

On citing what a search finds, Reactome says: It should be remembered that while we strive to contain the most current and accurate data, Reactome should not be used in citations where other primary sources of information are available. [R12] The record card prints the literature Reactome attaches to a curated pathway for that reason. The data are public domain: All data in the Reactome database and files derived from that data are licensed under the Creative Commons Public Domain Dedication (CC0). User may copy, modify, and distribute these data, even for commercial purposes, without asking for permission. Attribution is encouraged but not required. [R10] The pathway illustrations are licensed apart, under CC BY 4.0, and none is shown here.

A gene on this list is one Reactome places in this pathway in this release, and that is all the row says: it does not say the gene is expressed in a tissue or associated with a disease, which are the other two explorers' questions, read from other sources. Reactome's papers of record are [R01] [R02].

[R01] Ragueneau E, Gong C, Sinquin P, Sevilla C, Beavers D, Grentner A, et al. (2026). The Reactome Knowledgebase 2026. Nucleic Acids Research 54:D673-D681. PMID 41251150, doi 10.1093/nar/gkaf1223.
[R02] Milacic M, Beavers D, Conley P, Gong C, Gillespie M, Griss J, et al. (2024). The Reactome Pathway Knowledgebase 2024. Nucleic Acids Research 52:D672-D678. PMID 37941124, doi 10.1093/nar/gkad1025.
[R10] Reactome, reactome.org. License Agreement. https://reactome.org/license, read 2026-09-09.
[R12] Reactome, reactome.org. Citing us. https://reactome.org/cite, read 2026-09-09.
[R13] Reactome, reactome.org. Download. https://reactome.org/download-data, read 2026-09-09.

Stable id: R-HSA-446203.8
Class: Pathway
Species: Homo sapiens (human)
Inferred: no: a curated human pathway
Disease pathway: no
Released: 2010-06-15
Last updated: 2018-12-13
Events inside: 6, counted from the record

N-linked glycosylation is the most important form of post-translational modification for proteins synthesized and folded in the Endoplasmic Reticulum (Stanley et al. 2009). An early study in 1999 revealed that about 50% of the proteins in the Swiss-Prot database at the time were N-glycosylated (Apweiler et al. 1999). It is now established that the majority of the proteins in the secretory pathway require glycosylation in order to achieve proper folding.

The addition of an N-glycan to a protein can have several roles (Shental-Bechor & Levy 2009). First, glycans enhance the solubility and stability of the proteins in the ER, the golgi and on the outside of the cell membrane, where the composition of the medium is strongly hydrophilic and where proteins, that are mostly hydrophobic, have difficulty folding properly. Second, N-glycans are used as signal molecules during the folding and transport process of the protein: they have the role of labels to determine when a protein must interact with a chaperon, be transported to the golgi, or targeted for degradation in case of major folding defects. Third, and most importantly, N-glycans on completely folded proteins are involved in a wide range of processes: they help determine the specificity of membrane receptors in innate immunity or in cell-to-cell interactions, they can change the properties of hormones and secreted proteins, or of the proteins in the vesicular system inside the cell.

All N-linked glycans are derived from a common 14-sugar oligosaccharide synthesized in the ER, which is attached co-translationally to a protein while this is being translated inside the reticulum. The process of the synthesis of this glycan, known as Synthesis of the N-glycan precursor or LLO, constitutes one of the most conserved pathways in eukaryotes, and has been also observed in some eubacteria. The attachment usually happens on an asparagine residue within the consensus sequence asparagine-X-threonine by an complex called oligosaccharyl transferase (OST).

After being attached to an unfolded protein, the glycan is used as a label molecule in the folding process (also known as Calnexin/Calreticulin cycle) (Lederkremer 2009). The majority of the glycoproteins in the ER require at least one glycosylated residue in order to achieve proper folding, even if it has been shown that a smaller portion of the proteins in the ER can be folded without this modification.

Once the glycoprotein has achieved proper folding, it is transported via the cis-Golgi through all the Golgi compartments, where the glycan is further modified according to the properties of the glycoprotein. This process involves relatively few enzymes but due to its combinatorial nature, can lead to several millions of different possible modifications. The exact topography of this network of reactions has not been established yet, representing one of the major challenges after the sequencing of the human genome (Hossler et al. 2006).

Since N-glycosylation is involved in an great number of different processes, from cell-cell interaction to folding control, mutations in one of the genes involved in glycan assembly and/or modification can lead to severe development problems (often affecting the central nervous system). All the diseases in genes involved in glycosylation are collectively known as Congenital Disorders of Glycosylation (CDG) (Sparks et al. 2003), and classified as CDG type I for the genes in the LLO synthesis pathway, and CDG type II for the others.

Literature Reactome attaches

N-Glycans2009
Congenital Disorders of Glycosylation Overview1993
On the frequency of protein glycosylation, as deduced from analysis of the SWISS-PROT databaseBiochim Biophys Acta, 1999, vol. 1473, p. 4-8PubMed 10580125
Folding of glycoproteins: toward understanding the biophysics of the glycosylation codeCurr Opin Struct Biol, 2009, vol. 19, p. 524-33PubMed 19647993
Glycoprotein folding, quality control and ER-associated degradationCurr Opin Struct Biol, 2009, vol. 19, p. 515-23PubMed 19616933
GlycoVis: visualizing glycan distribution in the protein N-glycosylation pathway in mammalian cellsBiotechnol Bioeng, 2006, vol. 95, p. 946-60PubMed 16807922

Reactome's record lists 14 orthologous events, R-MMU-446203 (Mus musculus), R-RNO-446203 (Rattus norvegicus) among the species this site serves, and 12 in species this site opens no page for.

Reactome ContentService, this pathway's record · Reactome v97 · read 2026-09-30 ET · Reactome pathway browser"Asparagine N-linked glycosylation". Reactome, release 97, https://reactome.org/PathwayBrowser/#/R-HSA-446203 with StableID: R-HSA-446203.8 (accessed 2026-09-30 ET). Reactome version 97 (CC0).

Reading this pathway's record from Reactome (the pathway record).Still reading. A first read of a pathway can take a while; this page waits up to 35 seconds for it, and its scripts then bring in the panel, or a line saying what did not arrive.

02The genes

Genes Reactome places in this human pathway

The mapping files place 312 genes in this human pathway; showing 1 to 100, in pages of 100, sorted by symbol for reading. The order carries no ranking.

Genes Reactome places in this human pathway, page 1 of 4
Gene	Authority id	Mapping file id	Evidence codes
GeneACTR10	AuthorityHGNC:17372	Mapping file id55860 NCBI file	EvidenceTAS
GeneACTR1A	AuthorityHGNC:167	Mapping file id10121 NCBI file	EvidenceTAS
GeneALG1	AuthorityHGNC:18294	Mapping file id56052 NCBI file	EvidenceTAS
GeneALG10	AuthorityHGNC:23162	Mapping file id84920 NCBI file	EvidenceIEA
GeneALG10B	AuthorityHGNC:31088	Mapping file id144245 NCBI file	EvidenceIEA
GeneALG11	AuthorityHGNC:32456	Mapping file id440138 NCBI file	EvidenceIEA
GeneALG12	AuthorityHGNC:19358	Mapping file id79087 NCBI file	EvidenceTAS
GeneALG13	AuthorityHGNC:30881	Mapping file id79868 NCBI file	EvidenceTAS
GeneALG14	AuthorityHGNC:28287	Mapping file id199857 NCBI file	EvidenceTAS
GeneALG2	AuthorityHGNC:23159	Mapping file id85365 NCBI file	EvidenceIEA, TAS
GeneALG3	AuthorityHGNC:23056	Mapping file id10195 NCBI file	EvidenceTAS
GeneALG5	AuthorityHGNC:20266	Mapping file id29880 NCBI file	EvidenceTAS
GeneALG6	AuthorityHGNC:23157	Mapping file id29929 NCBI file	EvidenceTAS
GeneALG8	AuthorityHGNC:23161	Mapping file id79053 NCBI file	EvidenceTAS
GeneALG9	AuthorityHGNC:15672	Mapping file id79796 NCBI file	EvidenceTAS
GeneAMDHD2	AuthorityHGNC:24262	Mapping file id51005 NCBI file	EvidenceTAS
GeneAMFR	AuthorityHGNC:463	Mapping file id267 NCBI file	EvidenceTAS
GeneANK1	AuthorityHGNC:492	Mapping file id286 NCBI file	EvidenceTAS
GeneANK2	AuthorityHGNC:493	Mapping file id287 NCBI file	EvidenceTAS
GeneANK3	AuthorityHGNC:494	Mapping file id288 NCBI file	EvidenceTAS
GeneANKRD28	AuthorityHGNC:29024	Mapping file id23243 NCBI file	EvidenceTAS
GeneARCN1	AuthorityHGNC:649	Mapping file id372 NCBI file	EvidenceTAS
GeneAREG	AuthorityHGNC:651	Mapping file id374 NCBI file	EvidenceTAS
GeneARF1	AuthorityHGNC:652	Mapping file id375 NCBI file	EvidenceTAS
GeneARF3	AuthorityHGNC:654	Mapping file id377 NCBI file	EvidenceTAS
GeneARF4	AuthorityHGNC:655	Mapping file id378 NCBI file	EvidenceTAS
GeneARF5	AuthorityHGNC:658	Mapping file id381 NCBI file	EvidenceTAS
GeneARFGAP1	AuthorityHGNC:15852	Mapping file id55738 NCBI file	EvidenceTAS
GeneARFGAP2	AuthorityHGNC:13504	Mapping file id84364 NCBI file	EvidenceTAS
GeneARFGAP3	AuthorityHGNC:661	Mapping file id26286 NCBI file	EvidenceTAS
GeneASGR1	AuthorityHGNC:742	Mapping file id432 NCBI file	EvidenceTAS
GeneASGR2	AuthorityHGNC:743	Mapping file id433 NCBI file	EvidenceTAS
GeneB4GALNT2	AuthorityHGNC:24136	Mapping file id124872 NCBI file	EvidenceTAS
GeneB4GALT1	AuthorityHGNC:924	Mapping file id2683 NCBI file	EvidenceTAS
GeneB4GALT2	AuthorityHGNC:925	Mapping file id8704 NCBI file	EvidenceTAS
GeneB4GALT3	AuthorityHGNC:926	Mapping file id8703 NCBI file	EvidenceTAS
GeneB4GALT4	AuthorityHGNC:927	Mapping file id8702 NCBI file	EvidenceTAS
GeneB4GALT5	AuthorityHGNC:928	Mapping file id9334 NCBI file	EvidenceTAS
GeneB4GALT6	AuthorityHGNC:929	Mapping file id9331 NCBI file	EvidenceTAS
GeneBET1	AuthorityHGNC:14562	Mapping file id10282 NCBI file	EvidenceTAS
GeneBET1L	AuthorityHGNC:19348	Mapping file id51272 NCBI file	EvidenceTAS
GeneCALR	AuthorityHGNC:1455	Mapping file id811 NCBI file	EvidenceTAS
GeneCANX	AuthorityHGNC:1473	Mapping file id821 NCBI file	EvidenceTAS
GeneCAPZA1	AuthorityHGNC:1488	Mapping file id829 NCBI file	EvidenceTAS
GeneCAPZA2	AuthorityHGNC:1490	Mapping file id830 NCBI file	EvidenceTAS
GeneCAPZA3	AuthorityHGNC:24205	Mapping file id93661 NCBI file	EvidenceTAS
GeneCAPZB	AuthorityHGNC:1491	Mapping file id832 NCBI file	EvidenceTAS
GeneCD55	AuthorityHGNC:2665	Mapping file id1604 NCBI file	EvidenceTAS
GeneCD59	AuthorityHGNC:1689	Mapping file id966 NCBI file	EvidenceTAS
GeneCGA	AuthorityHGNC:1885	Mapping file id1081 NCBI file	EvidenceTAS
GeneCHST10	AuthorityHGNC:19650	Mapping file id9486 NCBI file	EvidenceTAS
GeneCHST8	AuthorityHGNC:15993	Mapping file id64377 NCBI file	EvidenceTAS
GeneCMAS	AuthorityHGNC:18290	Mapping file id55907 NCBI file	EvidenceTAS
GeneCNIH1	AuthorityHGNC:19431	Mapping file id10175 NCBI file	EvidenceTAS
GeneCNIH2	AuthorityHGNC:28744	Mapping file id254263 NCBI file	EvidenceTAS
GeneCNIH3	AuthorityHGNC:26802	Mapping file id149111 NCBI file	EvidenceTAS
GeneCOG1	AuthorityHGNC:6545	Mapping file id9382 NCBI file	EvidenceTAS
GeneCOG2	AuthorityHGNC:6546	Mapping file id22796 NCBI file	EvidenceTAS
GeneCOG3	AuthorityHGNC:18619	Mapping file id83548 NCBI file	EvidenceTAS
GeneCOG4	AuthorityHGNC:18620	Mapping file id25839 NCBI file	EvidenceTAS
GeneCOG5	AuthorityHGNC:14857	Mapping file id10466 NCBI file	EvidenceTAS
GeneCOG6	AuthorityHGNC:18621	Mapping file id57511 NCBI file	EvidenceTAS
GeneCOG7	AuthorityHGNC:18622	Mapping file id91949 NCBI file	EvidenceTAS
GeneCOG8	AuthorityHGNC:18623	Mapping file id84342 NCBI file	EvidenceTAS
GeneCOL7A1	AuthorityHGNC:2214	Mapping file id1294 NCBI file	EvidenceTAS
GeneCOPA	AuthorityHGNC:2230	Mapping file id1314 NCBI file	EvidenceTAS
GeneCOPB1	AuthorityHGNC:2231	Mapping file id1315 NCBI file	EvidenceTAS
GeneCOPB2	AuthorityHGNC:2232	Mapping file id9276 NCBI file	EvidenceTAS
GeneCOPE	AuthorityHGNC:2234	Mapping file id11316 NCBI file	EvidenceTAS
GeneCOPG1	AuthorityHGNC:2236	Mapping file id22820 NCBI file	EvidenceTAS
GeneCOPG2	AuthorityHGNC:2237	Mapping file id26958 NCBI file	EvidenceTAS
GeneCOPZ1	AuthorityHGNC:2243	Mapping file id22818 NCBI file	EvidenceTAS
GeneCOPZ2	AuthorityHGNC:19356	Mapping file id51226 NCBI file	EvidenceTAS
GeneCSNK1D	AuthorityHGNC:2452	Mapping file id1453 NCBI file	EvidenceTAS
GeneCTSA	AuthorityHGNC:9251	Mapping file id5476 NCBI file	EvidenceTAS
GeneCTSC	AuthorityHGNC:2528	Mapping file id1075 NCBI file	EvidenceTAS
GeneCTSZ	AuthorityHGNC:2547	Mapping file id1522 NCBI file	EvidenceTAS
GeneDAD1	AuthorityHGNC:2664	Mapping file id1603 NCBI file	EvidenceTAS
GeneDCTN1	AuthorityHGNC:2711	Mapping file id1639 NCBI file	EvidenceTAS
GeneDCTN2	AuthorityHGNC:2712	Mapping file id10540 NCBI file	EvidenceTAS
GeneDCTN3	AuthorityHGNC:2713	Mapping file id11258 NCBI file	EvidenceTAS
GeneDCTN4	AuthorityHGNC:15518	Mapping file id51164 NCBI file	EvidenceTAS
GeneDCTN5	AuthorityHGNC:24594	Mapping file id84516 NCBI file	EvidenceTAS
GeneDCTN6	AuthorityHGNC:16964	Mapping file id10671 NCBI file	EvidenceTAS
GeneDDOST	AuthorityHGNC:2728	Mapping file id1650 NCBI file	EvidenceTAS
GeneDERL1	AuthorityHGNC:28454	Mapping file id79139 NCBI file	EvidenceTAS
GeneDERL2	AuthorityHGNC:17943	Mapping file id51009 NCBI file	EvidenceTAS
GeneDHDDS	AuthorityHGNC:20603	Mapping file id79947 NCBI file	EvidenceTAS
GeneDHRSX	AuthorityHGNC:18399	Mapping file id207063 NCBI file	EvidenceTAS
GeneDOLK	AuthorityHGNC:23406	Mapping file id22845 NCBI file	EvidenceTAS
GeneDOLPP1	AuthorityHGNC:29565	Mapping file id57171 NCBI file	EvidenceTAS
GeneDPAGT1	AuthorityHGNC:2995	Mapping file id1798 NCBI file	EvidenceTAS
GeneDPM1	AuthorityHGNC:3005	Mapping file id8813 NCBI file	EvidenceTAS
GeneDPM2	AuthorityHGNC:3006	Mapping file id8818 NCBI file	EvidenceTAS
GeneDPM3	AuthorityHGNC:3007	Mapping file id54344 NCBI file	EvidenceTAS
GeneDYNC1H1	AuthorityHGNC:2961	Mapping file id1778 NCBI file	EvidenceTAS
GeneDYNC1I1	AuthorityHGNC:2963	Mapping file id1780 NCBI file	EvidenceTAS
GeneDYNC1I2	AuthorityHGNC:2964	Mapping file id1781 NCBI file	EvidenceTAS
GeneDYNC1LI1	AuthorityHGNC:18745	Mapping file id51143 NCBI file	EvidenceTAS
GeneDYNC1LI2	AuthorityHGNC:2966	Mapping file id1783 NCBI file	EvidenceTAS

Evidence codes on this page: IEA, TAS, as the mapping file writes them; a row carrying two was written twice by the file, once under each. The mapping file id says which file placed the gene: an NCBI Gene id from NCBI2Reactome_All_Levels.txt, an Ensembl gene id from Ensembl2Reactome_All_Levels.txt. The two files can disagree about a code, so a row's codes are the one file's view. Measured by this site's build over this release on 2026-09-09: of the 138,908 human pathway-gene pairs both files place, 207 (0.149 per cent, over 47 genes) carry TAS in the Ensembl file where the NCBI rows carry IEA alone.

Reactome mapping files, the human rows for this pathway · Reactome release 97 · read 2026-09-09 · Reactome downloads"NCBI2Reactome_All_Levels.txt" and "Ensembl2Reactome_All_Levels.txt", Reactome, release 97, https://reactome.org/download-data/ (date of access 2026-09-09). Data CC0; attribution encouraged.

03The hierarchy

Parents and children in this release's hierarchy

Parents

Post-translational protein modificationR-HSA-5975921,507 genes

Children

Reactome's hierarchy is a graph rather than a tree: a pathway can sit under more than one parent, and the gene counts are each pathway's own placements at every level under it, so a parent's count is not the sum of its children's.

Reactome, the human pathway list and hierarchy relationship file · Reactome release 97 · read 2026-09-09 · Reactome downloads"ReactomePathways.txt" and "ReactomePathwaysRelation.txt", Reactome, release 97, https://reactome.org/download-data/ (date of access 2026-09-09). Data CC0; attribution encouraged.