CODSWALLOP

Spike glycoprotein

Severe acute respiratory syndrome coronavirus 2 · seed P0DTC2 · 1273 aa · family defined as ≥30% identity to that seed · compiled 07 October 2026

CATH and SCOP identifiers come from the RCSB's own structure annotations, which the Domains panel already reads, so these are looked up rather than guessed at.

1,991Entries 2,000Entities 821Constructs 29Organisms 1,417Ligand-bound
0.96 ÅBest res.
3.20 ÅMedian res.

Every figure here is counted over the whole family rather than quoted from one entry.

The reference structure

7EAN, the structure every other member of this family is superposed onto. Rendered by the RCSB and embedded here: the live app shows an interactive viewport, which a document that fetches nothing cannot.

Rendered structure of 7EAN
7EAN at the RCSB · open it in the 3D viewer

Which residues anyone has ever seen

How many of this family's constructs contain each residue of the seed. A trough is a stretch nobody has put in a construct, which is a construct-design answer rather than a disorder one.

163612731949 constructs

Constructs, most-used first

821 distinct constructs across 1,991 entries. 1,968 polymer entities differ from the UniProt canonical sequence in some way, 601 carry a recognised expression tag and 5 carry a fusion partner.

"Differs from canonical" is not the same as "engineered". The canonical sequence is the full gene product, so a secreted protein whose structures all start after its signal peptide counts every one of them as different: lysozyme's most-used construct, residues 19–147 on 1,239 entities, is simply the mature protein. Read the construct column below for what was actually done, rather than this count.

EntitiesLengthBest (Å)Best entryWhat was made
56 1288 2.20 7LXY Strep-II; 1 internal deletion; 3-residue insertion after 1208; R682G, R683S, R685S +41 more
40 1288 2.62 7B18 Strep-II; 1 internal deletion; 3-residue insertion after 1208; R682G, R683S, R685S +37 more
36 205 1.71 8CWU residues 333-537; T531G, N532H, L533H +4 more
29 1283 2.70 7CZT FLAG; K986P, V987P
25 223 1.91 7EAN residues 319-541
24 205 1.85 7MZI residues 331-535; K528G, K529S, S530H +5 more
24 231 1.71 7JMP residues 319-549; N542S, F543G, N544H +5 more
20 195 1.75 7OLZ residues 333-527
17 194 2.10 7Y3O residues 334-527
17 1256 2.80 7UAP 3 internal deletions; 7-residue insertion after 1259; R685A, F817P, A892P +30 more
17 1286 3.16 7TOV Strep-II; 2 internal deletions; 3-residue insertion after 1208; T19R, G142D, R158G +43 more
16 194 2.40 7X2K residues 333-526
16 205 1.77 7NEH residues 324-528; S325T, I326G, V327H +6 more
16 1261 2.70 7DF3 FLAG+His9; residues 1-1224; 14-residue insertion after 1208; R682G, R683S, R685S +14 more
15 1280 2.80 7RU1 residues 1-1267; 1 internal deletion; 14-residue insertion after 1208; R682G, R683S, R685S +43 more
15 1288 2.80 7ND9 Strep-II; 1 internal deletion; 3-residue insertion after 1208; R682G, R683S, R685S +37 more
15 1288 3.20 7YVK 1 internal deletion; 14-residue insertion after 1208; T19I, G142D, V213G +83 more
14 1208 3.10 7YC5 residues 1-1208; R682G, R683S, R685S +6 more
14 1258 3.30 7WD0 FLAG+His9; residues 1-1224; 1 internal deletion; 14-residue insertion after 1208; L18F, D80A, D215G +23 more
14 1273 2.30 8CXQ K986P, V987P
13 202 2.30 8QRG residues 327-528; V327H, R328H, F329H +6 more
13 1281 2.70 7V8A 1 internal deletion; 6-residue insertion after 1227; T19R, G142D, R158G +64 more
12 1234 2.10 8G71 residues 14-1241; 6-residue insertion after 1227; D614G, R682A, R683G +31 more
12 1278 3.10 7X8W 1 internal deletion; 6-residue insertion after 1227; R682G, R683S, R685S +59 more
12 1281 2.60 7K43 His6; 2 internal deletions; 3-residue insertion after 12; M1S, F2L, V3L +49 more

Showing the 25 most-used of 821.

Positions people deliberately mutate

Columns where the wild-type residue still dominates but a real minority carries something else, which is a different question from "what varies across species".

K1211P 69% D614G 47% A942P 47% A892P 45% A899P 44% F817P 43% N501Y 34% E484A 33% K417N 32% I1210V 30% T478K 30% Q498R 26% G142D 25% P681H 25% S477N 25% T19I 24% S373P 23% Y505H 22% N440K 22% G339D 22% S371F 22% S375F 21% H655Y 19% N679K 19% V213G 18% D796Y 18% L452R 18% Q954H 18% N969K 18% D405N 18%

What it assembles into

Oligomeric stateChainsEntriesShare
trimeric3 843 42.3%
dimeric2 308 15.5%
pentameric5 211 10.6%
nonameric9 201 10.1%
tetrameric4 132 6.6%
hexameric6 126 6.3%
heptameric7 89 4.5%
pentadecameric15 20 1.0%

458 entries have the depositor's assembly corroborated by PISA, 1,526 carry the depositor's word alone and 7 were assigned by PISA where the depositor gave none. The middle figure is not a disagreement: PISA may have returned nothing or never run. 4 entries carry more than one assembly with different chain counts, so they have no single answer to quote: 3D0H, 3D0I, 8ELP, 9JTD.

Domain architecture

Every source's own domains on the seed axis, one row each. They are not merged: Pfam, CATH, SCOP and InterPro disagree about boundaries, and a merged track would state a consensus none of them gave.

CATHSpike glycoprotein, N-termImmunoglobulinsSCOP2BMetalloproteases (zincins)Concanavalin A-like lectinSARS coronavirus receptor-Coronavirus S1 glycoproteiCoronavirus S2 glycoproteiCoronavirus spike S2 glyco16361273
DomainSourceSpan (seed)Chains
Spike glycoprotein, N-terminal domainCATH 2.60.120.960 28–317 495
ImmunoglobulinsCATH 2.60.40.10 336–439 183
Metalloproteases (zincins), catalytic domainSCOP2B 8037105 15–610 31
Concanavalin A-like lectins/glucanasesSCOP2B 8092469 27–293 612
SARS coronavirus receptor-binding domain-likeSCOP2B 8092467 335–526 1,275
Coronavirus S1 glycoprotein, domain C-likeSCOP2B 8092661 529–592 852
Coronavirus S1 glycoprotein, domain D-likeSCOP2B 8092663 593–699 169
Coronavirus S2 glycoprotein stalk-likeSCOP2B 8092471 717–1068 850
Coronavirus spike S2 glycoprotein connector domain-likeSCOP2B 8092473 1074–1146 854

What binds it

NAG NAG1401 entries EIC EIC26 entries BLA BLA17 entries MAN MAN10 entries NDG NDG3 entries PG0 PG03 entries PRO PRO3 entries BLR BLR3 entries LEU LEU2 entries FOL FOL2 entries FUC FUC2 entries VCG VCG2 entries
ComponentClassNameEntriesBest (Å)
NAGcofactor 2-Acetamido-2-Deoxy-Beta-D-Glucopyranose 1,401 1.21
ZNion Zinc Ion 158 2.08
GOLcryoprotectant Glycerol 91 0.96
CLion Chloride Ion 83 1.55
SO4ion Sulfate Ion 49 1.70
EDOcryoprotectant 1,2-Ethanediol 47 1.21
EICligand Linoleic Acid 26 1.88
PEGcryoprotectant Di(Hydroxyethyl)ether 25 1.77
PO4ion Phosphate Ion 17 1.94
BLAligand Biliverdine Ix Alpha 17 1.82
CITbuffer Citric Acid 14 1.55
ACTcryoprotectant Acetate Ion 11 1.21
PG4cryoprotectant Tetraethylene Glycol 11 1.82
NAion Sodium Ion 11 1.55
MANcofactor Alpha-D-Mannopyranose 10 2.36
TRSbuffer 2-Amino-2-Hydroxymethyl-Propane-1,3-Diol 10 1.67
PGEcryoprotectant Triethylene Glycol 9 1.82
MGion Magnesium Ion 5 1.87
NO3ion Nitrate Ion 5 1.59
DMScryoprotectant Dimethyl Sulfoxide 4 2.42

How it crystallises

Parsed from the free text 545 depositors typed into _exptl_crystal_grow.pdbx_details, out of 545 entries that recorded anything at all. Median pH 7.0 (range 3.3 to 10.5).

Precipitants

PEG × Sodium citrate × Ammonium sulfate × Sodium chloride × Isopropanol × Lithium sulfate × Magnesium chloride × MPD × Sodium malonate × Tacsimate × Jeffamine × PEG (unspecified) × Sodium formate × Dioxane ×

Buffers

Citrate × Tris × HEPES × MES × Sodium acetate × Bis-Tris × Imidazole × Phosphate × Bis-Tris propane × Sodium cacodylate × CAPS × CHES ×

Which entries to trust

1,986 entries carry a wwPDB validation report: 1,465 clean, 399 worth a check and 122 with something to explain. Median clashscore 6.02, median RSRZ outliers 3.72%, median R-free minus R-work 0.041. 1,984 have released structure factors.

Across species

OrganismEntriesBest (Å)Ligand-boundSeed covered
Severe acute respiratory syndrome coronavirus 21,832 0.96 1307 100%
Severe acute respiratory syndrome coronavirus32 2.40 19 100%
Severe acute respiratory syndrome-related coronavirus28 2.20 14 96%
Homo sapiens18 1.90 14 95%
Bat coronavirus RaTG1312 2.60 11 95%
Sarbecovirus11 2.13 10 95%
Bat SARS-like coronavirus WIV19 1.88 8 95%
Pangolin coronavirus9 2.45 6 17%
Human betacoronavirus 2c EMC/20125 2.80 2 84%
Middle East respiratory syndrome-related coronavirus4 2.59 3 83%
Saccharomyces cerevisiae S288C4 2.70 4 21%
Bat SARS-like coronavirus RsSHC0143 2.24 2 95%

Seed sequence

1273 residues, numbered every ten. Every identity figure in this document is measured against this sequence.

active or binding site modified residue or glycosylation disulphide cysteine transmembrane or signal the 15 most-substituted positions

1MFVFLVLLPL VSSQCVNLTT RTQLPPAYTN SFTRGVYYPD KVFRSSVLHS TQDLFLPFFS
61NVTWFHAIHV SGTNGTKRFD NPVLPFNDGV YFASTEKSNI IRGWIFGTTL DSKTQSLLIV
121NNATNVVIKV CEFQFCNDPF LGVYYHKNNK SWMESEFRVY SSANNCTFEY VSQPFLMDLE
181GKQGNFKNLR EFVFKNIDGY FKIYSKHTPI NLVRDLPQGF SALEPLVDLP IGINITRFQT
241LLALHRSYLT PGDSSSGWTA GAAAYYVGYL QPRTFLLKYN ENGTITDAVD CALDPLSETK
301CTLKSFTVEK GIYQTSNFRV QPTESIVRFP NITNLCPFGE VFNATRFASV YAWNRKRISN
361CVADYSVLYN SASFSTFKCY GVSPTKLNDL CFTNVYADSF VIRGDEVRQI APGQTGKIAD
421YNYKLPDDFT GCVIAWNSNN LDSKVGGNYN YLYRLFRKSN LKPFERDIST EIYQAGSTPC
481NGVEGFNCYF PLQSYGFQPT NGVGYQPYRV VVLSFELLHA PATVCGPKKS TNLVKNKCVN
541FNFNGLTGTG VLTESNKKFL PFQQFGRDIA DTTDAVRDPQ TLEILDITPC SFGGVSVITP
601GTNTSNQVAV LYQDVNCTEV PVAIHADQLT PTWRVYSTGS NVFQTRAGCL IGAEHVNNSY
661ECDIPIGAGI CASYQTQTNS PRRARSVASQ SIIAYTMSLG AENSVAYSNN SIAIPTNFTI
721SVTTEILPVS MTKTSVDCTM YICGDSTECS NLLLQYGSFC TQLNRALTGI AVEQDKNTQE
781VFAQVKQIYK TPPIKDFGGF NFSQILPDPS KPSKRSFIED LLFNKVTLAD AGFIKQYGDC
841LGDIAARDLI CAQKFNGLTV LPPLLTDEMI AQYTSALLAG TITSGWTFGA GAALQIPFAM
901QMAYRFNGIG VTQNVLYENQ KLIANQFNSA IGKIQDSLSS TASALGKLQD VVNQNAQALN
961TLVKQLSSNF GAISSVLNDI LSRLDKVEAE VQIDRLITGR LQSLQTYVTQ QLIRAAEIRA
1021SANLAATKMS ECVLGQSKRV DFCGKGYHLM SFPQSAPHGV VFLHVTYVPA QEKNFTTAPA
1081ICHDGKAHFP REGVFVSNGT HWFVTQRNFY EPQIITTDNT FVSGNCDVVI GIVNNTVYDP
1141LQPELDSFKE ELDKYFKNHT SPDVDLGDIS GINASVVNIQ KEIDRLNEVA KNLNESLIDL
1201QELGKYEQYI KWPWYIWLGF IAGLIAIVMV TIMLCCMTSC CSCLKGCCSC GSCCKFDEDD
1261SEPVLKGVKL HYT

Sites are UniProt's curated features where the seed is a UniProt accession; the substituted positions are measured from this family's own alignment rather than annotated, and only the fifteen most substituted are marked: every position carrying a minority substitution would be most of the protein, because the family holds orthologues. A residue can carry more than one and is drawn with the first that applies, in the order of the key above.

Primary citations

One record per paper, not per entry.

YearCitation
2026 Monoclonal antibodies from COVID-19 convalescent patients target cryptic epitopes for broad SARS-CoV-2 neutralization. Proc.Natl.Acad.Sci.USA doi:10.1073/pnas.2523864123
2026 Mining antibody functionality via AI-guided structural landscape profiling. Nat Commun doi:10.1038/s41467-026-70553-6
2026 Modification of a SARS-CoV-2 spike-RBD targeting nanobody for pH-dependent binding and its chromatographic engineering for purification of WT and variant S-RBDs with mild elution conditions. Int.J.Biol.Macromol. doi:10.1016/j.ijbiomac.2025.149749
2026 IgG-Bridging-Seeded Synergistic Aggregation of SARS-CoV-2 Spikes Underlies Potent Neutralization by a Low-Affinity Antibody. Adv Sci doi:10.1002/advs.202517192
2026 A structural and mechanistic atlas of NTD antibody neutralization and immune escape across SARS-CoV-2 prototype and its (sub-)variants. Proc.Natl.Acad.Sci.USA doi:10.1073/pnas.2535385123
2026 In vivo evolution of antibody CR3022 expands cross-neutralization of SARS-CoV-2 variants and informs pan-sarbecovirus immunity. Cell Rep doi:10.1016/j.celrep.2026.117137
2026 Somatic Evolution of a Germline Antibody Expands its Breadth to Neutralize Early SARS-CoV-2 Omicron Variants. Adv Sci doi:10.1002/advs.76522
2026 mRNA delivery of a class 1/4 SARS-CoV-2 neutralizing antibody protects against diverse sarbecoviruses in a lethal mouse challenge model. Proc.Natl.Acad.Sci.USA doi:10.1073/pnas.2536870123
2026 Epitope-focused discovery of SARS-CoV-2 antibodies that potently neutralize Omicron variants. Nat Microbiol doi:10.1038/s41564-026-02282-x
2026 TMPRSS2-mediated coronavirus spike activation and inhibition. Nat.Struct.Mol.Biol. doi:10.1038/s41594-026-01801-y
2026 Intranasal administration of broad-spectrum macrocyclic peptide inhibitor protects against SARS-CoV-2 Omicron variants. Nat Commun doi:10.1038/s41467-026-68462-9
2026 Defining the mechanism of cross-reactivity for a SARS-CoV-2 Beta-elicited antibody toward omicron sub-lineages. Structure doi:10.1016/j.str.2026.01.006
2026 The buried S2 apex of SARS-CoV-2 spike elicits an immunodominant germline-restricted public antibody response. Biorxiv doi:10.64898/2026.02.18.706653
2026 Virological characteristics of SARS-CoV-2-related coronaviruses dynamically circulating in Southeast Asia. Cell doi:10.1016/j.cell.2026.04.019
2026 Twenty-year persistence of SARS-CoV-1 immune imprinting shapes antibody responses to SARS-CoV-2 infection. Immunity doi:10.1016/j.immuni.2026.08.009
2026 Enteric alpha-defensins contribute to intestinal mucosal immunity against SARS-CoV-2 infection. Mucosal Immunol doi:10.1016/j.mucimm.2026.100392
2026 Broad Neutralizing Activity of Monoclonal Antibodies Against the Omicron Variants Isolated From Patients With Early Severe Acute Respiratory Syndrome Coronavirus-2. J Med Virol doi:10.1002/jmv.70969
2026 Mass spectrometry-based mapping of conformational epitopes on SARS-CoV-2 antigens targeted by monoclonal antibodies. Int.J.Biol.Macromol. doi:10.1016/j.ijbiomac.2026.153102
2026 Structural and functional characterization of a conserved cryptic epitope on SARS-CoV-2 spike S2 subunit. Plos Pathog. doi:10.1371/journal.ppat.1014391
2026 Biophysical trade-offs in antibody evolution are resolved by conformation-mediated epistasis. Biorxiv doi:10.64898/2026.03.12.711465
2026 The computationally designed TRI2-2 miniprotein inhibitor protects against multiple SARS-CoV-2 Omicron variants. Commun Biol doi:10.1038/s42003-025-09499-2
2026 Species- and variant-specific ACE2 compatibility shapes SARS-CoV-2 spillover potential in North American cervids. Nat Commun doi:10.1038/s41467-026-71623-5
2026 Cryo-EM structure of locked spike glycoprotein from bat SARS-like coronavirus WIV1, molecular dynamics and biophysics across host range. Proc.Natl.Acad.Sci.USA doi:10.1073/pnas.2516874123
2026 Recurrent SARS-CoV-2 Omicron broadly neutralizing humanized antibodies in different single human V H 1-2-rearranging mouse models. Proc.Natl.Acad.Sci.USA doi:10.1073/pnas.2537053123
2026 Neutralization of SARS-CoV-2 by IgM-14 via engagement of two distinct spike epitopes. Plos Pathog. doi:10.1371/journal.ppat.1014071
2026 Molecular basis of fox ACE2 recognition by receptor binding domains of SARS-CoV-2 and PCoV-GD. Cell Insight doi:10.1016/j.cellin.2026.100314
2026 Structure-Guided Design of Therapeutic Antibodies Targeting SARS-CoV-2 Omicron Variants. Res Sq doi:10.21203/rs.3.rs-9917568/v1
2026 Public antibody clonotypes and deep learning identify SARS-CoV-2 and HIV broadly neutralizing antibodies in immune repertoires. Cell Rep doi:10.1016/j.celrep.2026.117582
2026 Steric hindrance of antibody binding in an Omicron spike fusion intermediate. Nature doi:10.1038/s41586-026-10462-2
2026 Development of a Thermostable and Broadly Neutralizing Pan-Sarbecovirus Vaccine Candidate. Acs Infect Dis. doi:10.1021/acsinfecdis.5c00479
2025 Integrating immune library probing with structure-based computational design to develop potent neutralizing nanobodies against emerging SARS-CoV-2 variants. Mabs doi:10.1080/19420862.2025.2499595
2025 Ultra-potent RBM-specific single-domain antibody broadly neutralizes multiple SARS-CoV-2 variants with picomolar activity. Int.J.Biol.Macromol. doi:10.1016/j.ijbiomac.2025.145386
2025 Structure-guided engineering of a mutation-tolerant inhibitor peptide against variable SARS-CoV-2 spikes. Proc.Natl.Acad.Sci.USA doi:10.1073/pnas.2413465122
2025 Broadly neutralizing antibodies targeting a conserved silent face of spike RBD resist extreme SARS-CoV-2 antigenic drift Cell Rep doi:10.1016/j.celrep.2025.115948
2025 Affinity maturation endows potent activity onto class 6 SARS-CoV-2 broadly neutralizing antibodies. Proc.Natl.Acad.Sci.USA doi:10.1073/pnas.2417544121
2025 Rapid discovery of monoclonal antibodies by microfluidics-enabled FACS of single pathogen-specific antibody-secreting cells. Nat.Biotechnol. doi:10.1038/s41587-024-02346-5
2025 Production and cryo-electron microscopy structure of an internally tagged SARS-CoV-2 spike ecto-domain construct. J Struct Biol X doi:10.1016/j.yjsbx.2025.100123
2025 Engineering a multivalent antibody nanoparticle to overcome SARS-CoV-2 Omicron immune evasion. Plos Pathog. doi:10.1371/journal.ppat.1013744
2025 Mapping of human monoclonal antibody responses to XBB.1.5 COVID-19 monovalent vaccines: a B cell analysis. Lancet Microbe doi:10.1016/j.lanmic.2025.101103
2025 Cross-reactive sarbecovirus antibodies induced by mosaic RBD nanoparticles. Proc.Natl.Acad.Sci.USA doi:10.1073/pnas.2501637122