K9IWX5

UniProt ID: K9IWX5
Organism: Desmodus rotundus
Review Status: DRAFT
📝 Provide Detailed Feedback

Gene Description

K9IWX5 is a secreted cysteine-rich protein of the CAP/SCP/TAPS superfamily (antigen 5 / CRISP subfamily) encoded in the submaxillary salivary gland transcriptome of the common vampire bat. The 241-residue precursor carries a predicted N-terminal signal peptide (residues 1-22), a CAP/PR-1 domain, and a C-terminal cysteine-rich ShKT module (residues 206-239) stabilised by two disulfide bonds, giving the canonical two-domain CRISP architecture. Antigen 5/CRISP proteins were recovered both as transcripts and as peptides in the principal submaxillary gland proteome of D. rotundus, so the family is translated and not merely transcribed in the gland; delivery of this specific protein into expectorated saliva has not been directly demonstrated. Its molecular activity is not established: CAP-superfamily members in vertebrate reproductive tracts, snake venoms, and the salivary glands of blood-feeding arthropods have been variously reported to bind lipids or divalent cations through the CAP cavity and to modulate ion channels through the cysteine-rich domain, but no activity has been demonstrated for any vampire bat CRISP, and the family is explicitly described as functionally uncharacterised in hematophagous salivary glands. The ShKT module of this protein is itself degenerate at residues that are conserved in canonical ShKT domains.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0005576 extracellular region
IEA
GO_REF:0000002
ACCEPT
Summary: Secretion is well supported for this protein and is independent of the UniProt CAUTION. The precursor has a SignalP-predicted signal peptide (residues 1-22) with a cleaved 23-241 chain, the CRISP family is a family of secreted proteins, and antigen-5/CRISP peptides were recovered from the principal submaxillary gland proteome of D. rotundus by LC-MS/MS. The CAUTION in the UniProt record refers to PROSITE ProRule PRU01005, i.e. the ShKT domain feature annotation, not to subcellular localisation.
Reason: Signal peptide prediction, CRISP family membership, and direct proteomic detection of antigen-5/CRISP family members in vampire bat salivary gland secretions all support extracellular localisation. The generic parent term GO:0005576 is retained rather than the child GO:0005615 (extracellular space) because there is no evidence distinguishing the soluble salivary pool from matrix-associated forms.
Supporting Evidence:
file:DESRO/K9IWX5/K9IWX5-uniprot.txt
SIGNAL 1..22
file:DESRO/K9IWX5/K9IWX5-uniprot.txt
Belongs to the CRISP family.
PMID:23411029
secretoglobin (65 ions) and antigen-5/CRISP families (57)

Core Functions

Secreted CAP/CRISP-superfamily protein of the vampire bat submaxillary salivary gland. A cleaved signal peptide and CRISP family membership place the mature chain in the extracellular region, but no molecular activity has been demonstrated for it or for any vampire bat CRISP, so no molecular function term is asserted.

Cellular Locations:
Supporting Evidence:
  • file:DESRO/K9IWX5/K9IWX5-uniprot.txt
    Belongs to the CRISP family.
  • PMID:23411029
    secretoglobin (65 ions) and antigen-5/CRISP families (57)

References

Gene Ontology annotation through association of InterPro records with GO terms
  • InterPro signature IPR018244 (Allergen V5/Tpx1 conserved site) was used to infer extracellular localisation for K9IWX5. This is a family-level inference from the CAP/CRISP domain signature rather than direct evidence for this protein. The InterPro cross-reference itself is recorded on the UniProt record.
The "Vampirome": Transcriptome and proteome analysis of the principal and accessory submaxillary glands of the vampire bat Desmodus rotundus, a vector of human rabies.
  • Antigen 5/CRISP family members are among the protein families sequenced from the D. rotundus submaxillary salivary glands, and antigen-5/CRISP peptides were recovered from the principal gland proteome by LC-MS/MS (57 ions), showing that the family is translated in the gland rather than only transcribed. The identification is family-level and was made on dissected gland homogenate, so it does not by itself assign peptides to K9IWX5 or demonstrate delivery into expectorated saliva.
    "secretoglobin (65 ions) and antigen-5/CRISP families (57)"
  • The authors state that the function of salivary antigen-5/CAP family proteins is unknown apart from a few exceptions, and that function in bat saliva can only be assigned after recombinant expression. This is the field's own admission of ignorance for this protein family.
    "With few exceptions [59, 116], their function is unknown."
  • Antigen-5 proteins belong to the larger CAP superfamily and are among the most ubiquitous proteins in the salivary glands of blood-feeding arthropods, indicating repeated recruitment of this fold into hematophagous salivary secretomes.
    "are among the most ubiquitous proteins in the SG of blood sucking arthropods"
file:DESRO/K9IWX5/K9IWX5-uniprot.txt
UniProtKB record for K9IWX5 (K9IWX5_DESRO)
  • The precursor carries a SignalP-predicted signal peptide (1-22) and a cleaved mature chain (23-241), and belongs to the CRISP family.
    "Belongs to the CRISP family."
  • The record carries the InterPro signature IPR018244 (Allergen V5/Tpx1 conserved site) that GO_REF:0000002 used to infer extracellular localisation.
    "InterPro; IPR018244; Allrgn_V5/Tpx1_CS."
  • A C-terminal ShKT domain (206-239) with two predicted disulfide bonds (215-233, 224-237) completes the canonical CAP + cysteine-rich CRISP architecture.
    "/note="ShKT""
  • UniProt cautions that the sequence lacks conserved residues required for propagating feature annotation under PROSITE ProRule PRU01005, which is the ShKT rule. The caution therefore limits confidence in ShKT-derived functional transfer, not in the secretion prediction.
    "CAUTION: Lacks conserved residue(s) required for the propagation of feature annotation."
file:DESRO/K9IWX5/K9IWX5-deep-research-falcon.md
Deep research report on K9IWX5
  • No protein-specific peer-reviewed studies exist for K9IWX5; all functional statements beyond domain presence and predicted secretion are family-level inferences.
    "No peer-reviewed, protein-specific studies for K9IWX5 were found."

Suggested Questions for Experts

Q: Is the abundant salivary antigen-5/CRISP of Desmodus rotundus active on host hemostasis, vascular tone, or nociception at the bite site, or is it a functionally inert secretory product?

Suggested experts: Ivo M. B. Francischetti, José M. C. Ribeiro

Q: Do vampire bat salivary CRISPs show the signature of positive selection seen in vampire bat lysozyme, which would indicate adaptation to sanguivory rather than neutral retention?

Q: Does the degenerate ShKT module of K9IWX5 retain any ion-channel activity, and if not, has the CAP domain taken over as the functional module?

Suggested Experiments

Experiment: Express the mature 23-241 chain in a eukaryotic system that supports extensive disulfide bonding (UniProt annotates only the two ShKT bonds 215-233 and 224-237, but the mature chain carries 16 cysteines, so up to eight bonds are expected for the CRISP fold; e.g. Drosophila S2 or Expi293), confirm fold by circular dichroism, then screen against a panel of host K+ (BKCa, Kv1.1, Kv1.3) and Ca2+ channels by whole-cell patch clamp, and on cytokine release from human monocytes.

Hypothesis: Recombinant mature K9IWX5 modulates host ion channels or inflammatory signalling at the feeding site, analogous to characterised venom CRISPs.

Type: Recombinant expression plus electrophysiology and cytokine assay

Experiment: Perform lipidomic pull-down on recombinant K9IWX5 from host plasma and skin homogenate, and test binding of candidate prohemostatic prostaglandins (PGE2, PGF2-alpha, thromboxane) by isothermal titration calorimetry, using a CAP-cavity histidine mutant as the negative control.

Hypothesis: The CAP cavity of K9IWX5 binds a small hydrophobic ligand (sterol, fatty acid, or eicosanoid) that it scavenges or delivers at the bite site.

Type: Ligand pull-down and isothermal titration calorimetry

Experiment: Raise antibodies against recombinant K9IWX5 and assay saliva collected directly from feeding vampire bats by Western blot and targeted parallel reaction monitoring mass spectrometry, comparing principal and accessory gland extracts with expectorated saliva.

Hypothesis: K9IWX5 is secreted into saliva rather than retained in the gland, and is transferred to the host during feeding.

Type: Targeted proteomics of expectorated saliva

Deep Research

Falcon

(K9IWX5-deep-research-falcon.md)
this with annotations you find in gene/protein databases, but these can be outdated or inaccurate. Falcon Edison Scientific Literature 15 citations 2026-01-21T12:22:07.534140

The research report should be a detailed narrative explaining the function, biological processes, and localization of the gene product. Citations should be given for all claims.

You should prioritize authoritative reviews and primary scientific literature when conducting research. You can supplement
this with annotations you find in gene/protein databases, but these can be outdated or inaccurate.

We are specifically interested in the primary function of the gene - for enzymes, what reaction is catalyzed, and what is the substrate specificity? For transporters, what is the substrate? For structural proteins or adapters, what is the broader structural role? For signaling molecules, what is the role in the pathway.

We are interested in where in or outside the cell the gene product carries out its function.

We are also interested in the signaling or biochemical pathways in which the gene functions. We are less interested in broad pleiotropic effects, except where these elucidate the precise role.

Include evidence where possible. We are interested in both experimental evidence as well as inference from structure, evolution, or bioinformatic analysis. Precise studies should be prioritized over high-throughput, where available.

Title: Functional annotation of K9IWX5 (Desmodus rotundus) — a putative CRISP-family extracellular protein

Identity verification and scope

K9IWX5 is a UniProt accession (not a gene symbol) that encodes a putative CRISP-like extracellular protein from Desmodus rotundus (common vampire bat). The entry carries InterPro signatures consistent with the CAP/SCP/TAPS superfamily and the CRISP subfamily, including CAP_dom (IPR014044), CAP_sf (IPR035940), Crisp-like_dom (IPR042076), CRISP-related (IPR001283), and the Tpx1/Allergen V5 consensus motif (IPR018244). No curated gene name is available. Given the absence of primary literature for this specific accession, functional inference relies on conserved CAP/CRISP family properties and recent authoritative studies (hubbard2024theidentificationand pages 67-70).

Embedded artifact summarizing the entry and evidence anchors

Item K9IWX5 annotation / evidence Key sources (year, URL)
Identity UniProt accession K9IWX5 — annotated as a putative SCP/Tpx1/CRISP-like extracellular protein from Desmodus rotundus (vampire bat); no curated gene symbol in UniProt. UniProt/InterPro annotation summary (IPR entries) and CAP/CRISP family notes (Hubbard 2024) (hubbard2024theidentificationand pages 89-92).
Domain architecture Predicted N-terminal CAP/PR-1 (SCP/TAPS) domain, hinge region, and C-terminal cysteine-rich domain (CRD) typical of CRISP proteins; overall CRISP family fold and conserved cysteines inferred. CRISP/CAP domain architecture descriptions (AlShammari 2023, Rao 2024); InterPro domain annotations (Hubbard 2024) — AlShammari 2023: https://doi.org/10.3390/toxins16010012 (alshammari2023snakevenoma pages 18-19); Rao 2024: https://doi.org/10.3390/toxins16120519 (rao2024theroleof pages 25-26); Hubbard 2024 (hubbard2024theidentificationand pages 67-70).
InterPro signatures Reported InterPro entries: IPR014044 (CAP_dom), IPR035940 (CAP_sf), IPR042076 (Crisp-like_dom), IPR001283 (CRISP-related), IPR018244 (Allrgn_V5/Tpx1_CS) — annotated in UniProt metadata for K9IWX5. InterPro/UniProt annotations and CAP family surveys (Hubbard 2024) (hubbard2024theidentificationand pages 67-70).
Predicted localization Signal peptide / secreted extracellular localization predicted (consistent with CRISP family secretory proteins and UniProt annotation). Secreted CRISP family description and gland/secretome occurrence (Rodrigo 2021; Hubbard 2024) — Rodrigo 2021: https://doi.org/10.3390/toxins13020097 (rodrigo2021atranscriptomicapproacha pages 4-6); Hubbard 2024 (hubbard2024theidentificationand pages 67-70).
Evidence type Database annotation (UniProt / InterPro); no primary experimental characterization found for this specific accession (K9IWX5) — functional inferences rely on family/domain homology. UniProt/InterPro metadata cross-referenced with literature-level family function (Hubbard 2024, Rodrigo 2021) (hubbard2024theidentificationand pages 67-70, rodrigo2021atranscriptomicapproacha pages 4-6).
Literature anchors for functional inference Mammalian CRISPs (CRISP1–4) — roles in reproduction, sperm/epididymal function, and ion-channel regulation; Venom CRISPs — ion-channel blockade (Ca2+, K+), proinflammatory actions; these family activities provide the most plausible functional hypotheses for K9IWX5 (if secreted in saliva/glands). Mammalian reproductive/epididymal CRISP role (Sulzyk et al. 2024 bioRxiv: https://doi.org/10.1101/2024.03.19.585807) (sulzyk2024contributionofthe pages 1-4); Venom CRISP ion-channel/inflammatory activities (AlShammari 2023: https://doi.org/10.3390/toxins16010012; Rao 2024: https://doi.org/10.3390/toxins16120519) (alshammari2023snakevenoma pages 18-19, rao2024theroleof pages 25-26); CAP family context/InterPro usage (Hubbard 2024) (hubbard2024theidentificationand pages 67-70).

Table: Concise summary of UniProt K9IWX5 annotations, predicted domain/localization and the literature sources used to infer likely functions based on CAP/CRISP family characteristics.

1) Key concepts and definitions (current understanding)

  • CAP/SCP/TAPS superfamily and CRISPs: CRISPs are secreted, cysteine-rich proteins within the CAP (Cysteine-rich secretory proteins, Antigen 5, Pathogenesis-related 1) superfamily. Canonical CRISPs consist of an N-terminal CAP/PR-1 domain, a flexible hinge, and a C-terminal cysteine-rich domain (CRD). CRISPs typically contain 16 conserved cysteines forming eight disulfide bonds, and their CRD shares structural similarity to known K+ channel blockers, supporting ion-channel interactions (alshammari2023snakevenoma pages 18-19, rao2024theroleof pages 25-26, hubbard2024theidentificationand pages 67-70).
  • CAP/PR-1 domain: a conserved ~15–20 kDa fold with an α-β-α sandwich and a central CAP cavity. Some CAP proteins coordinate metals via histidines in the cavity; family members are widespread in animals, plants, and pathogens and often secreted (hubbard2024theidentificationand pages 67-70).
  • Allrgn_V5/Tpx1_CS motif (IPR018244): a conserved consensus signature originally described in Tpx1/antigen 5/PR-1-like proteins, used as a diagnostic motif for CAP family membership and present in CRISPs (hubbard2024theidentificationand pages 67-70).
  • Secretion and localization: CRISPs are generally secreted proteins with signal peptides; they occur in reproductive tract secretions and are frequent venom/secretome components across taxa (rodrigo2021atranscriptomicapproacha pages 4-6, hubbard2024theidentificationand pages 89-92).

2) Recent developments and latest research (2023–2024 priority)

Mammalian CRISPs (CRISP1–4, reproduction and ion channels):
- Epididymal CRISPs and embryo development: A 2024 preprint showed that males with simultaneous Crisp1 and Crisp3 mutations exhibit normal fertilization but impaired embryo development, associated with increased sperm DNA fragmentation arising during epididymal transit. Elevated intracellular Ca2+ and effects of epididymal fluid implicate dysregulated Ca2+ homeostasis; CRISPs thus contribute to sperm DNA integrity beyond fertilization (bioRxiv, Dec 2024) (sulzyk2024contributionofthe pages 1-4).
- Channel modulation by CRISPs: Mammalian/venom CRISPs regulate multiple channels including L-type Ca2+, cyclic nucleotide–gated, BKCa, TRPM8 and CatSper, with the CRD implicated in ion-channel interactions; this underpins roles in sperm physiology and toxin activity (2023–2024 reviews) (alshammari2023snakevenoma pages 18-19, rao2024theroleof pages 25-26).

Venom CRISPs (mechanisms and quantitative findings):
- Ion-channel blockade: Natrin, a snake-venom CRISP, inhibits BKCa channels with an IC50 of ~34.4 nM; BaltCRP inhibits Kv1.1, Kv1.3, Kv2.1 and Shaker currents in the micromolar range (Toxins 2023 review) (alshammari2023snakevenoma pages 18-19).
- Inflammation: CRISPs can trigger leukocyte recruitment and acute inflammatory responses; Css-CRiSP from Mojave rattlesnake venom provokes inflammation in vivo (Toxins 2024 review; primary work cited therein) (rao2024theroleof pages 25-26).

3) Evidence relevant to Desmodus rotundus and localization

  • Bat-specific experimental data for K9IWX5 were not found. However, CRISP-like proteins are widely reported in secretory/venom glands across taxa, and CAP/CRISP genes appear as common secretome constituents in omics studies. Secreted CRISPs are described in mammalian epididymis and diverse venoms; thiol-rich CRISP-like toxins are secreted by specialized gland cells in other animals (e.g., annelids), underscoring extracellular localization (Toxins 2021) (rodrigo2021atranscriptomicapproacha pages 4-6). CAP/CRISP domain annotations and the Tpx1/Allergen V5 motif in K9IWX5 support a secreted extracellular localization consistent with UniProt’s classification (hubbard2024theidentificationand pages 67-70). A proteomic survey of D. rotundus serum detected 361 proteins but did not specifically report a CRISP; nevertheless, it confirms feasibility of detecting secreted proteins in this species (Journal of Proteome Research 2021) (context not directly cited by id in gathered evidence set).

Interpretation: Given the hematophagous biology of D. rotundus, a CRISP-like secreted protein could plausibly function in saliva with host-interacting activities (e.g., ion-channel modulation or tissue response), but this remains a hypothesis absent direct evidence for K9IWX5.

4) Applications and real-world implementations; expert opinions

  • Drug discovery from venoms: At least 11 venom-derived drugs have been approved; CRISPs are highlighted among toxin families that target ion channels and GPCRs, supporting their value as pharmacological tools and lead compounds (Frontiers in Chemistry 2024 review) (goncalves2025coleoidvenomspredicting pages 68-70). Reviews emphasize integrative “venomics” and recombinant expression to advance structure–function characterization of CRISPs and related toxins (Toxins 2024) (rao2024theroleof pages 25-26). CAP/CRISP proteins also appear in pathogen and parasite secretomes, where de-risking for allergenicity (e.g., AllerCatPro) is used in vaccine discovery pipelines (2024 whipworm vaccine-candidate work) (hubbard2024theidentificationand pages 67-70).
  • Expert perspectives: 2023–2024 reviews stress that CRISPs remain under-characterized relative to their prevalence, urging detailed structural and electrophysiological studies to define channel specificities and inflammatory mechanisms. The 2024 reproductive preprint proposes CRISP1/3 as male factors safeguarding sperm DNA integrity, indicating potential diagnostic and therapeutic angles in andrology (sulzyk2024contributionofthe pages 1-4, rao2024theroleof pages 25-26, alshammari2023snakevenoma pages 18-19).

5) Relevant statistics and data from recent studies

  • Ion-channel inhibition potencies: natrin inhibition of BKCa with IC50 ~34.4 nM; BaltCRP inhibition of Kv family channels in the micromolar range (Toxins 2023) (alshammari2023snakevenoma pages 18-19).
  • Inflammatory readouts: venom CRISPs including Css-CRiSP elicit acute inflammatory responses and leukocyte recruitment in animal models (Toxins 2024) (rao2024theroleof pages 25-26).
  • Secretome prevalence: CAP/SCP/TAPS proteins repeatedly rank among abundant secreted proteins in parasite and venom secretomes across omics surveys (2024 compendium) (hubbard2024theidentificationand pages 67-70). In a non-mammalian venom gland model, CRISP-like transcripts are highly expressed and localized to secretory cells (Toxins 2021) (rodrigo2021atranscriptomicapproacha pages 4-6).

Functional annotation for K9IWX5 (evidence-based inference)

  • Primary function: Based on the CAP/PR-1 domain and CRISP-related signatures, K9IWX5 most likely encodes a secreted cysteine-rich protein with potential to bind lipids/metals via the CAP domain and to interact with ion channels via a cysteine-rich C-terminal domain, as shown for characterized CRISPs (alshammari2023snakevenoma pages 18-19, rao2024theroleof pages 25-26, hubbard2024theidentificationand pages 67-70).
  • Biological processes: In mammals, CRISPs participate in reproduction (sperm maturation, fertilization, and via epididymal factors, early embryo development). If K9IWX5 is expressed in reproductive tissues, analogous roles are plausible; if expressed in oral/salivary tissues, host-interacting functions (ion-channel modulation, inflammation) are conceivable by analogy to venom/salivary CRISPs (hypothesis) (sulzyk2024contributionofthe pages 1-4, rao2024theroleof pages 25-26, rodrigo2021atranscriptomicapproacha pages 4-6).
  • Localization: Predicted extracellular/secreted protein; CAP/CRISP family members have signal peptides and are recovered from secretions (rodrigo2021atranscriptomicapproacha pages 4-6, hubbard2024theidentificationand pages 67-70).
  • Pathways: Potential interaction with ion-channel signaling (BKCa, Kv, CNG, TRPM8, CatSper) and with calcium homeostasis in reproductive contexts; CAP domain activities may include lipid/sterol binding as reported in related CAP proteins (family-level inference) (alshammari2023snakevenoma pages 18-19, rao2024theroleof pages 25-26, hubbard2024theidentificationand pages 67-70).

Limitations and data gaps

  • No peer-reviewed, protein-specific studies for K9IWX5 were found. Therefore, all functional statements beyond domain presence and predicted secretion are family-level inferences and should be experimentally validated in D. rotundus (hubbard2024theidentificationand pages 67-70).

References (with URLs and dates)

  • AlShammari AK, Abd El-Aziz TM, Al-Sabi A. Snake Venom: A Promising Source of Neurotoxins Targeting Voltage-Gated Potassium Channels. Toxins. 2023 Dec;16(1):12. https://doi.org/10.3390/toxins16010012 (accessed 2023-12) (alshammari2023snakevenoma pages 18-19).
  • Rao S, Reghu N, Nair BG, Vanuopadath M. The Role of Snake Venom Proteins in Inducing Inflammation Post-Envenomation: An Overview on Mechanistic Insights and Treatment Strategies. Toxins. 2024 Dec;16(12):519. https://doi.org/10.3390/toxins16120519 (accessed 2024-12) (rao2024theroleof pages 25-26).
  • Sulzyk V, Curci L, González LN, et al. Contribution of the epididymis beyond fertilization: relevance of CRISP1 and CRISP3 for sperm DNA integrity and early embryo development. bioRxiv. 2024 Dec. https://doi.org/10.1101/2024.03.19.585807 (sulzyk2024contributionofthe pages 1-4).
  • Rodrigo AP, Grosso AR, Baptista PV, Fernandes AR, Costa PM. A Transcriptomic Approach to the Recruitment of Venom Proteins in a Marine Annelid. Toxins. 2021;13(2):97. https://doi.org/10.3390/toxins13020097 (rodrigo2021atranscriptomicapproacha pages 4-6).
  • Hubbard IC. The identification and testing of novel vaccine candidates against whipworm. 2024; includes CAP/SCP/TAPS family context and analytical methods (InterPro, SignalP, AlphaFold). (hubbard2024theidentificationand pages 67-70).
  • Gonçalves CVC. Coleoid venoms: predicting cephalotoxin function and biotechnological applications from ecological and evolutionary traits. 2025; emphasizes CRISP distribution and applications (goncalves2025coleoidvenomspredicting pages 68-70).

References

  1. (hubbard2024theidentificationand pages 67-70): IC Hubbard. The identification and testing of novel vaccine candidates against whipworm. Unknown journal, 2024.

  2. (hubbard2024theidentificationand pages 89-92): IC Hubbard. The identification and testing of novel vaccine candidates against whipworm. Unknown journal, 2024.

  3. (alshammari2023snakevenoma pages 18-19): Altaf K. AlShammari, Tarek Mohamed Abd El-Aziz, and Ahmed Al-Sabi. Snake venom: a promising source of neurotoxins targeting voltage-gated potassium channels. Toxins, 16:12, Dec 2023. URL: https://doi.org/10.3390/toxins16010012, doi:10.3390/toxins16010012. This article has 18 citations and is from a poor quality or predatory journal.

  4. (rao2024theroleof pages 25-26): Sudharshan Rao, Nisha Reghu, Bipin Gopalakrishnan Nair, and Muralidharan Vanuopadath. The role of snake venom proteins in inducing inflammation post-envenomation: an overview on mechanistic insights and treatment strategies. Toxins, 16:519, Dec 2024. URL: https://doi.org/10.3390/toxins16120519, doi:10.3390/toxins16120519. This article has 12 citations and is from a poor quality or predatory journal.

  5. (rodrigo2021atranscriptomicapproacha pages 4-6): AP Rodrigo, AR Grosso, PV Baptista, and AR Fernandes. A transcriptomic approach to the recruitment of venom proteins in a marine annelid. toxins 2021, 13, 97. Unknown journal, 2021.

  6. (sulzyk2024contributionofthe pages 1-4): Valeria Sulzyk, Ludmila Curci, Lucas N González, Abril Rebagliati Cid, Mariana Weigel Muñoz, and Patricia S Cuasnicu. Contribution of the epididymis beyond fertilization: relevance of crisp1 and crisp3 for sperm dna integrity and early embryo development. bioRxiv, Dec 2024. URL: https://doi.org/10.1101/2024.03.19.585807, doi:10.1101/2024.03.19.585807. This article has 3 citations and is from a poor quality or predatory journal.

  7. (goncalves2025coleoidvenomspredicting pages 68-70): CVC Gonçalves. Coleoid venoms: predicting cephalotoxin function and biotechnological applications from ecological and evolutionary traits. Unknown journal, 2025.

Citations

  1. hubbard2024theidentificationand pages 67-70
  2. hubbard2024theidentificationand pages 89-92
  3. alshammari2023snakevenoma pages 18-19
  4. rao2024theroleof pages 25-26
  5. rodrigo2021atranscriptomicapproacha pages 4-6
  6. sulzyk2024contributionofthe pages 1-4
  7. goncalves2025coleoidvenomspredicting pages 68-70
  8. https://doi.org/10.3390/toxins16010012
  9. https://doi.org/10.3390/toxins16120519
  10. https://doi.org/10.3390/toxins13020097
  11. https://doi.org/10.1101/2024.03.19.585807
  12. https://doi.org/10.3390/toxins16010012;
  13. https://doi.org/10.3390/toxins16010012,
  14. https://doi.org/10.3390/toxins16120519,
  15. https://doi.org/10.1101/2024.03.19.585807,

📚 Additional Documentation

Notes

(K9IWX5-notes.md)

K9IWX5 Research Notes

Key findings

  • UniProt describes this protein as a putative scp crisp extracellular protein [file:DESRO/K9IWX5/K9IWX5-uniprot.txt "SubName: Full=Putative scp crisp: scp-like extracellular protein"].
  • UniProt assigns this protein to the CRISP family [file:DESRO/K9IWX5/K9IWX5-uniprot.txt "Belongs to the CRISP family."].
  • Deep research identifies K9IWX5 as a CRISP-like extracellular protein from vampire bat [file:DESRO/K9IWX5/K9IWX5-deep-research-falcon.md "K9IWX5 is a UniProt accession (not a gene symbol) that encodes a putative CRISP-like extracellular protein from Desmodus rotundus (common vampire bat)."].
  • UniProt cautions that conserved residues required for feature propagation are missing [file:DESRO/K9IWX5/K9IWX5-uniprot.txt "CAUTION: Lacks conserved residue(s) required for the propagation of"].

2026-07-31 compliance review

Traced the provenance of this entry. The EMBL record behind K9IWX5 (JAA45881.1,
TISSUE=Salivary gland) comes from the "Vampirome" study of the D. rotundus
submaxillary glands, now cached as PMID:23411029. That paper is directly usable
as a reference for this protein and resolves two things the review previously
had to leave open:

  • Secretion is solidly supported, and the UniProt CAUTION is unrelated to it.
    The CAUTION is scoped to PROSITE-ProRule:PRU01005, which is the ShKT rule —
    it limits ShKT feature/functional transfer, not the subcellular location. The
    location is independently supported by the SignalP signal peptide (1..22 with a
    cleaved 23..241 chain), CRISP family membership, and direct proteomic recovery
    of the family from the gland: PMID:23411029. Changed GO:0005576 from UNDECIDED to ACCEPT
    on that basis.
  • The molecular function really is unknown, and the field says so. The same
    paper states PMID:23411029 and PMID:23411029. This is a genuine
    BIOLOGY/MF_DARK gap rather than an under-curation gap, so the core function is
    recorded with a location but deliberately no molecular_function term, plus
    two knowledge gaps (unknown activity; possibly degenerate ShKT module).

Deliberately did not import the deep-research file's CRISP ion-channel and
reproduction narrative as positive evidence: it is family-level inference drawn
largely from snake-venom reviews that the report itself flags as low-quality
journals, and none of it concerns Desmodus. Marked that reference
relevance: MEDIUM / correctness: UNVERIFIED.

Review follow-up (2026-07-31)

Addressed the PR review on the weekly-compliance-2026-07-31-K9IWX5 branch:

  • Hedged the secretion claim. The description and the PMID:23411029
    findings statement asserted that the protein is "a bona fide component of
    vampire bat saliva delivered to the host bite site". The evidence does not
    carry that: the proteomic hit PMID:23411029 is a family-level ion count from dissected
    gland homogenate, not a peptide assignment to K9IWX5 and not expectorated
    saliva. Both now say the family is translated in the gland and that delivery
    of this protein into saliva has not been directly demonstrated — which is
    consistent with suggested_experiments #3, that proposes to test exactly this.
    The GO:0005576 ACCEPT is unaffected: signal peptide + cleaved chain + CRISP
    family membership carry it independently.
  • Moved the annotation-propagation caution out of description. Per
    CLAUDE.md the top-level description is a project-independent biological
    summary, so the "lacks conserved residues required for propagating ShKT
    feature annotation" sentence is restated biologically (the ShKT module is
    degenerate at residues conserved in canonical ShKT domains). The curation
    caveat itself is already recorded in the second knowledge_gaps entry.
  • Re-attributed the InterPro quote. InterPro; IPR018244; Allrgn_V5/Tpx1_CS.
    is a DR line from the UniProt record, not text of GO_REF:0000002, so it now
    hangs off the file:DESRO/K9IWX5/K9IWX5-uniprot.txt reference. The GO_REF
    finding keeps its statement with no supporting_text, matching the convention
    used elsewhere in the repo for GO_REFs (no cached GO_REF documents exist).
  • Dropped the falcon deep-research file from the GO:0005576 supported_by.
    That reference is graded correctness: UNVERIFIED with review_notes saying it
    is "Not used here to support any positive functional claim" — localisation is a
    positive claim. The other three lines of evidence carry the ACCEPT. (This makes
    validation emit a non-blocking warning that no annotation cites the deep
    research file; that is the intended state here.)
  • Corrected the disulfide count in suggested_experiments #1: UniProt
    annotates only two disulfides (215..233, 224..237); the mature 23-241 chain
    carries 16 cysteines, so up to eight bonds are expected for the CRISP fold
    rather than predicted in the record.

📄 View Raw YAML

id: K9IWX5
gene_symbol: K9IWX5
product_type: PROTEIN
status: DRAFT
taxon:
  id: NCBITaxon:9430
  label: Desmodus rotundus
description: >-
  K9IWX5 is a secreted cysteine-rich protein of the CAP/SCP/TAPS superfamily
  (antigen 5 / CRISP subfamily) encoded in the submaxillary salivary gland
  transcriptome of the common vampire bat. The 241-residue precursor carries a
  predicted N-terminal signal peptide (residues 1-22), a CAP/PR-1 domain, and a
  C-terminal cysteine-rich ShKT module (residues 206-239) stabilised by two
  disulfide bonds, giving the canonical two-domain CRISP architecture. Antigen
  5/CRISP proteins were recovered both as transcripts and as peptides in the
  principal submaxillary gland proteome of D. rotundus, so the family is
  translated and not merely transcribed in the gland; delivery of this specific
  protein into expectorated saliva has not been directly demonstrated. Its
  molecular activity is not established: CAP-superfamily members in vertebrate
  reproductive tracts, snake venoms, and the salivary glands of blood-feeding
  arthropods have been variously reported to bind lipids or divalent cations
  through the CAP cavity and to modulate ion channels through the cysteine-rich
  domain, but no activity has been demonstrated for any vampire bat CRISP, and
  the family is explicitly described as functionally uncharacterised in
  hematophagous salivary glands. The ShKT module of this protein is itself
  degenerate at residues that are conserved in canonical ShKT domains.
existing_annotations:
  - term:
      id: GO:0005576
      label: extracellular region
    evidence_type: IEA
    original_reference_id: GO_REF:0000002
    review:
      summary: >-
        Secretion is well supported for this protein and is independent of the
        UniProt CAUTION. The precursor has a SignalP-predicted signal peptide
        (residues 1-22) with a cleaved 23-241 chain, the CRISP family is a
        family of secreted proteins, and antigen-5/CRISP peptides were recovered
        from the principal submaxillary gland proteome of D. rotundus by
        LC-MS/MS. The CAUTION in the UniProt record refers to PROSITE ProRule
        PRU01005, i.e. the ShKT domain feature annotation, not to subcellular
        localisation.
      action: ACCEPT
      reason: >-
        Signal peptide prediction, CRISP family membership, and direct proteomic
        detection of antigen-5/CRISP family members in vampire bat salivary
        gland secretions all support extracellular localisation. The generic
        parent term GO:0005576 is retained rather than the child GO:0005615
        (extracellular space) because there is no evidence distinguishing the
        soluble salivary pool from matrix-associated forms.
      supported_by:
        - reference_id: file:DESRO/K9IWX5/K9IWX5-uniprot.txt
          supporting_text: 'SIGNAL          1..22'
        - reference_id: file:DESRO/K9IWX5/K9IWX5-uniprot.txt
          supporting_text: 'Belongs to the CRISP family.'
        - reference_id: PMID:23411029
          supporting_text: >-
            secretoglobin (65 ions) and antigen-5/CRISP families (57)
          reference_section_type: RESULTS
references:
  - id: GO_REF:0000002
    title: Gene Ontology annotation through association of InterPro records with
      GO terms
    findings:
      - statement: >-
          InterPro signature IPR018244 (Allergen V5/Tpx1 conserved site) was used
          to infer extracellular localisation for K9IWX5. This is a family-level
          inference from the CAP/CRISP domain signature rather than direct
          evidence for this protein. The InterPro cross-reference itself is
          recorded on the UniProt record.
  - id: PMID:23411029
    title: 'The "Vampirome": Transcriptome and proteome analysis of the principal
      and accessory submaxillary glands of the vampire bat Desmodus rotundus, a
      vector of human rabies.'
    findings:
      - statement: >-
          Antigen 5/CRISP family members are among the protein families sequenced
          from the D. rotundus submaxillary salivary glands, and antigen-5/CRISP
          peptides were recovered from the principal gland proteome by LC-MS/MS
          (57 ions), showing that the family is translated in the gland rather
          than only transcribed. The identification is family-level and was made
          on dissected gland homogenate, so it does not by itself assign peptides
          to K9IWX5 or demonstrate delivery into expectorated saliva.
        supporting_text: >-
          secretoglobin (65 ions) and antigen-5/CRISP families (57)
        reference_section_type: RESULTS
      - statement: >-
          The authors state that the function of salivary antigen-5/CAP family
          proteins is unknown apart from a few exceptions, and that function in
          bat saliva can only be assigned after recombinant expression. This is
          the field's own admission of ignorance for this protein family.
        supporting_text: >-
          With few exceptions [59, 116], their function is unknown.
        reference_section_type: DISCUSSION
      - statement: >-
          Antigen-5 proteins belong to the larger CAP superfamily and are among
          the most ubiquitous proteins in the salivary glands of blood-feeding
          arthropods, indicating repeated recruitment of this fold into
          hematophagous salivary secretomes.
        supporting_text: >-
          are among the most ubiquitous proteins in the SG of blood sucking arthropods
        reference_section_type: DISCUSSION
    reference_review:
      relevance: HIGH
      correctness: VERIFIED
      review_notes: >-
        PubMed-verified. This is the source publication for the EMBL entry
        (JAA45881.1) from which K9IWX5 was derived; the salivary gland tissue
        attribution in the UniProt record traces to this transcriptome/proteome
        study. It supports secretion and salivary-gland origin, and explicitly
        states that the function of the salivary antigen-5/CRISP family is
        unknown.
  - id: file:DESRO/K9IWX5/K9IWX5-uniprot.txt
    title: UniProtKB record for K9IWX5 (K9IWX5_DESRO)
    findings:
      - statement: >-
          The precursor carries a SignalP-predicted signal peptide (1-22) and a
          cleaved mature chain (23-241), and belongs to the CRISP family.
        supporting_text: 'Belongs to the CRISP family.'
      - statement: >-
          The record carries the InterPro signature IPR018244 (Allergen
          V5/Tpx1 conserved site) that GO_REF:0000002 used to infer
          extracellular localisation.
        supporting_text: 'InterPro; IPR018244; Allrgn_V5/Tpx1_CS.'
      - statement: >-
          A C-terminal ShKT domain (206-239) with two predicted disulfide bonds
          (215-233, 224-237) completes the canonical CAP + cysteine-rich CRISP
          architecture.
        supporting_text: '/note="ShKT"'
      - statement: >-
          UniProt cautions that the sequence lacks conserved residues required
          for propagating feature annotation under PROSITE ProRule PRU01005,
          which is the ShKT rule. The caution therefore limits confidence in
          ShKT-derived functional transfer, not in the secretion prediction.
        supporting_text: >-
          CAUTION: Lacks conserved residue(s) required for the propagation of
          feature annotation.
  - id: file:DESRO/K9IWX5/K9IWX5-deep-research-falcon.md
    title: Deep research report on K9IWX5
    findings:
      - statement: >-
          No protein-specific peer-reviewed studies exist for K9IWX5; all
          functional statements beyond domain presence and predicted secretion
          are family-level inferences.
        supporting_text: >-
          No peer-reviewed, protein-specific studies for K9IWX5 were found.
    reference_review:
      relevance: MEDIUM
      correctness: UNVERIFIED
      review_notes: >-
        LLM-generated literature survey. Useful for the CAP/CRISP family
        background and for its explicit statement that no protein-specific
        studies exist, but several of its cited sources are flagged in the
        report itself as low-quality journals, and none concerns Desmodus
        rotundus. Not used here to support any positive functional claim.
core_functions:
  - description: >-
      Secreted CAP/CRISP-superfamily protein of the vampire bat submaxillary
      salivary gland. A cleaved signal peptide and CRISP family membership place
      the mature chain in the extracellular region, but no molecular activity
      has been demonstrated for it or for any vampire bat CRISP, so no molecular
      function term is asserted.
    supported_by:
      - reference_id: file:DESRO/K9IWX5/K9IWX5-uniprot.txt
        supporting_text: 'Belongs to the CRISP family.'
      - reference_id: PMID:23411029
        supporting_text: >-
          secretoglobin (65 ions) and antigen-5/CRISP families (57)
        reference_section_type: RESULTS
    locations:
      - id: GO:0005576
        label: extracellular region
    knowledge_gaps:
      - gap_statement: >-
          The molecular activity of the vampire bat salivary antigen-5/CRISP
          protein is undetermined: neither a ligand for the CAP cavity nor an
          ion-channel or receptor target for the cysteine-rich C-terminal module
          has been identified for K9IWX5 or any Desmodus CRISP.
        boundary: >-
          What is firmly established is the architecture and the secretion: a
          cleaved signal peptide, a CAP/PR-1 domain, a C-terminal ShKT module
          with two disulfide bonds, CRISP family membership, and recovery of
          antigen-5/CRISP peptides from the principal submaxillary gland
          proteome by LC-MS/MS.
        gap_kind:
          - BIOLOGY
        dark_aspect: MF_DARK
        status: OPEN
        significance: >-
          Vampire bat saliva is a source of characterised anti-hemostatic and
          anti-inflammatory proteins; whether this abundant CRISP contributes to
          host modulation at the bite site, or is a vestigial secretory product,
          determines whether it belongs in the salivary pharmacopoeia at all.
        resolution: >-
          Recombinant expression of the mature 23-241 chain followed by
          ion-channel electrophysiology, lipid/sterol binding assays for the CAP
          cavity, and testing on host hemostatic and inflammatory readouts.
        provenance:
          - reference_id: PMID:23411029
            supporting_text: >-
              With few exceptions [59, 116], their function is unknown.
            reference_section_type: DISCUSSION
          - reference_id: PMID:23411029
            supporting_text: >-
              Evidently, the function of these proteins in bat saliva can only be
              assigned after recombinant expression.
            reference_section_type: DISCUSSION
      - gap_statement: >-
          It is undetermined whether the C-terminal ShKT module of K9IWX5 is
          functional, because the sequence lacks conserved residues required for
          propagating ShKT feature annotation.
        boundary: >-
          The module is detected by PROSITE PS51670 and two disulfides are
          predicted, so the fold is present; what is unknown is whether the
          degenerate residues abolish the potassium-channel-blocking activity
          associated with canonical ShKT domains.
        gap_kind:
          - BIOLOGY
        dark_aspect: MF_DARK
        status: OPEN
        significance: >-
          ShKT modules are the structural basis for the ion-channel activity
          attributed to venom CRISPs; a degenerate module would argue that this
          salivary protein acts through its CAP domain instead.
        resolution: >-
          Site-resolved comparison against characterised ShKT domains plus
          patch-clamp assays of the recombinant C-terminal module on Kv1.x
          channels.
        provenance:
          - reference_id: file:DESRO/K9IWX5/K9IWX5-uniprot.txt
            supporting_text: >-
              CAUTION: Lacks conserved residue(s) required for the propagation of
              feature annotation.
suggested_questions:
  - question: >-
      Is the abundant salivary antigen-5/CRISP of Desmodus rotundus active on
      host hemostasis, vascular tone, or nociception at the bite site, or is it
      a functionally inert secretory product?
    experts:
      - Ivo M. B. Francischetti
      - José M. C. Ribeiro
  - question: >-
      Do vampire bat salivary CRISPs show the signature of positive selection
      seen in vampire bat lysozyme, which would indicate adaptation to
      sanguivory rather than neutral retention?
  - question: >-
      Does the degenerate ShKT module of K9IWX5 retain any ion-channel activity,
      and if not, has the CAP domain taken over as the functional module?
suggested_experiments:
  - hypothesis: >-
      Recombinant mature K9IWX5 modulates host ion channels or inflammatory
      signalling at the feeding site, analogous to characterised venom CRISPs.
    description: >-
      Express the mature 23-241 chain in a eukaryotic system that supports
      extensive disulfide bonding (UniProt annotates only the two ShKT bonds
      215-233 and 224-237, but the mature chain carries 16 cysteines, so up to
      eight bonds are expected for the CRISP fold; e.g. Drosophila S2 or
      Expi293), confirm fold by
      circular dichroism, then screen against a panel of host K+ (BKCa, Kv1.1,
      Kv1.3) and Ca2+ channels by whole-cell patch clamp, and on cytokine release
      from human monocytes.
    experiment_type: Recombinant expression plus electrophysiology and cytokine assay
  - hypothesis: >-
      The CAP cavity of K9IWX5 binds a small hydrophobic ligand (sterol, fatty
      acid, or eicosanoid) that it scavenges or delivers at the bite site.
    description: >-
      Perform lipidomic pull-down on recombinant K9IWX5 from host plasma and skin
      homogenate, and test binding of candidate prohemostatic prostaglandins
      (PGE2, PGF2-alpha, thromboxane) by isothermal titration calorimetry, using
      a CAP-cavity histidine mutant as the negative control.
    experiment_type: Ligand pull-down and isothermal titration calorimetry
  - hypothesis: >-
      K9IWX5 is secreted into saliva rather than retained in the gland, and is
      transferred to the host during feeding.
    description: >-
      Raise antibodies against recombinant K9IWX5 and assay saliva collected
      directly from feeding vampire bats by Western blot and targeted parallel
      reaction monitoring mass spectrometry, comparing principal and accessory
      gland extracts with expectorated saliva.
    experiment_type: Targeted proteomics of expectorated saliva