dyf-6

UniProt ID: Q0G838
Organism: Caenorhabditis elegans
Review Status: COMPLETE
Aliases:
IFT46 F46F6.4
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Gene Description

dyf-6 encodes the Caenorhabditis elegans ortholog of intraflagellar transport protein 46 (IFT46), a structural subunit of intraflagellar transport (IFT) complex B. IFT-B, with IFT-A and the kinesin-2 and dynein-2 motors, drives the bidirectional movement of ciliary cargo that builds and maintains the cilia of ciliated sensory neurons. The 471-residue protein has a large N-terminal disordered/acidic region and no recognizable catalytic domain. DYF-6 is expressed in ciliated amphid and phasmid sensory neurons (and hypodermis), localizes to the cilium, the ciliary base/basal body region, dendrites and the neuronal cell body, and undergoes processive IFT movement within the ciliated dendritic endings. It is required to build full-length sensory cilia: in dyf-6 mutants the amphid and phasmid ciliary endings are foreshortened, the IFT-B marker OSM-6 is mislocalized in a pattern typical of complex B mutants, and the animals are defective in dye filling and chemotaxis. The ciliary role is conserved, with orthologs in Drosophila (CG15161, expressed in sensory cilia) and mammals (human IFT46). C. elegans has only non-motile sensory cilia, so the cilium DYF-6 builds is a non-motile sensory cilium.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0005815 microtubule organizing center
IBA
GO_REF:0000033
MODIFY
Summary: IFT46 orthologs concentrate at and inject IFT trains from the ciliary base. In C. elegans this is the basal body / transition-zone region; UniProt curates DYF-6 basal-body localization from PMID:16648645. The basal body is a type of microtubule organizing center, so the term is not wrong, but the specific "ciliary basal body" (GO:0036064) is the accurate, informative CC.
Reason: Term too general. DYF-6's documented location at the ciliary base is better captured by ciliary basal body (GO:0036064) than by the generic microtubule organizing center.
Propagation Review
Root cause: TERM SCOPING PROBLEM
Failure modes: GRANULARITY MISMATCH
Sources checked:
PANTHER:PTN000332669 · IFT46 family node SUPPORTS TRANSFER
The IFT46 family node correctly places DYF-6 at the microtubule organizing center (basal body), but the transferred term is coarser than the specific ciliary basal body location documented for the worm protein.
Proposed replacements: ciliary basal body
GO:0030992 intraciliary transport particle B
IBA
GO_REF:0000033
ACCEPT
Summary: DYF-6/IFT46 is a subunit of intraflagellar transport complex B (IFT-B). This phylogenetic transfer is corroborated by Bell et al.'s inference from the OSM-6 mislocalization pattern and by biochemical assignment to ComplexPortal CPX-1290.
Reason: Core cellular-component identity of DYF-6 as an IFT-B structural subunit.
Supporting Evidence:
PMID:28479320
intraflagellar transport (IFT)-B complex abolishes dynein-2's ciliary localization
file:worm/dyf-6/dyf-6-deep-research-falcon.md
IFT46 is a pivotal subunit of the IFT-B1 core subcomplex
GO:0060271 cilium assembly
IBA
GO_REF:0000033
ACCEPT
Summary: IFT is required to assemble and maintain cilia; dyf-6 mutants have foreshortened amphid and phasmid ciliary endings, so DYF-6 is required for cilium assembly.
Reason: Core biological process, supported experimentally in worm (PMID:16648645). The more specific worm-appropriate term non-motile cilium assembly is also annotated (IMP).
Supporting Evidence:
PMID:16648645
the cilia of the amphid and phasmid dendritic endings are foreshortened
GO:0031514 motile cilium
IBA
GO_REF:0000033
MARK AS OVER ANNOTATED
Summary: This IBA transfer from mammalian IFT46 assigns a motile-cilium location. C. elegans, however, has no motile cilia β€” all its cilia, including the amphid and phasmid sensory cilia in which DYF-6 acts, are non-motile. The matching worm annotation is non-motile cilium assembly (GO:1905515, IMP).
Reason: Taxonomically inappropriate over-propagation: C. elegans lacks motile cilia. DYF-6 acts in non-motile sensory cilia; the generic ciliary location (cilium, GO:0005929) is separately and correctly annotated.
Propagation Review
Root cause: PROPAGATION BAD
Failure modes: LINEAGE OR TAXON MISMATCH
Sources checked:
PANTHER:PTN000332669 · IFT46 family node SUPPORTS SOURCE BUT NOT TARGET
The IFT46 family node includes vertebrate members that function in motile cilia, but the C. elegans ortholog acts only in non-motile sensory cilia, so the motile-cilium location does not transfer.
GO:0042073 intraciliary transport
IBA
GO_REF:0000033
ACCEPT
Summary: DYF-6 functions in intraflagellar (intraciliary) transport as an IFT-B subunit; DYF-6::GFP undergoes IFT movement within ciliated endings.
Reason: Core biological process, confirmed experimentally in worm (PMID:16648645).
Supporting Evidence:
PMID:16648645
Movement of DYF-6::GFP within the ciliated endings of the neurons indicates that DYF-6 is involved in IFT
IEA
GO_REF:0000044
ACCEPT
Summary: Electronic mapping of the UniProt cilium subcellular-location keyword. DYF-6 is directly observed within cilia, where it undergoes IFT.
Reason: Correct core ciliary location, corroborated by experimental IFT movement within ciliated endings (PMID:16648645).
Supporting Evidence:
PMID:16648645
Movement of DYF-6::GFP within the ciliated endings of the neurons indicates that DYF-6 is involved in IFT
GO:0030425 dendrite
IEA
GO_REF:0000044
KEEP AS NON CORE
Summary: DYF-6 is present in the sensory dendrites through which IFT cargo is trafficked to the ciliated ending; UniProt curates dendrite localization from PMID:16648645 and WormBase makes the same call by IDA.
Reason: Real localization (the dendritic route to the cilium) but not the core site of DYF-6 function, which is the cilium/IFT machinery.
GO:0042073 intraciliary transport
IEA
GO_REF:0000002
ACCEPT
Summary: InterPro2GO transfer (IPR022088, IFT complex B) to intraciliary transport. Duplicates the IBA/NAS/IDA intraciliary-transport annotations and is correct.
Reason: Core biological process; redundant with experimental IDA (PMID:16648645) but consistent.
Supporting Evidence:
PMID:16648645
Movement of DYF-6::GFP within the ciliated endings of the neurons indicates that DYF-6 is involved in IFT
GO:0043204 perikaryon
IEA
GO_REF:0000044
KEEP AS NON CORE
Summary: Electronic mapping of the UniProt perikaryon subcellular-location keyword, the same call made experimentally (EXP) from PMID:16648645. DYF-6 is present in the neuronal cell body/perikaryon in addition to the cilium.
Reason: Real localization but not the core functional site; the cell body reflects the neuron of expression rather than DYF-6's IFT role.
GO:0120025 plasma membrane bounded cell projection
IEA
GO_REF:0000117
MARK AS OVER ANNOTATED
Summary: ARBA machine-learning annotation to a high-level grouping term that is the parent of both cilium and dendrite, which are already specifically annotated for DYF-6.
Reason: Over-general electronic grouping term; its informative descendants (cilium, dendrite) are already annotated, so it adds no information.
NAS
PMID:28479320
Dynein-Driven Retrograde Intraflagellar Transport Is Triphas...
ACCEPT
Summary: ComplexPortal (CPX-1290) assertion that DYF-6 localizes to the cilium, consistent with the experimental observation that DYF-6 undergoes IFT within cilia.
Reason: Correct core ciliary location.
Supporting Evidence:
PMID:16648645
Movement of DYF-6::GFP within the ciliated endings of the neurons indicates that DYF-6 is involved in IFT
GO:0030992 intraciliary transport particle B
NAS
PMID:28479320
Dynein-Driven Retrograde Intraflagellar Transport Is Triphas...
ACCEPT
Summary: ComplexPortal (CPX-1290) assignment of DYF-6 to IFT complex B, based on the affinity-purification / mass-spectrometry of the worm IFT-B complex in Yi et al. 2017 and reflected in the UniProt SUBUNIT statement.
Reason: Core cellular-component identity; biochemically supported IFT-B membership.
Supporting Evidence:
PMID:28479320
intraflagellar transport (IFT)-B complex abolishes dynein-2's ciliary localization
GO:0042073 intraciliary transport
NAS
PMID:28479320
Dynein-Driven Retrograde Intraflagellar Transport Is Triphas...
ACCEPT
Summary: ComplexPortal assertion that DYF-6, as an IFT-B subunit, functions in intraciliary transport, consistent with the worm experimental data.
Reason: Core biological process; consistent with IFT-B membership and experimental IFT movement (PMID:16648645).
Supporting Evidence:
PMID:16648645
Movement of DYF-6::GFP within the ciliated endings of the neurons indicates that DYF-6 is involved in IFT
GO:0060271 cilium assembly
NAS
PMID:28479320
Dynein-Driven Retrograde Intraflagellar Transport Is Triphas...
ACCEPT
Summary: ComplexPortal assertion that DYF-6 functions in cilium assembly, consistent with the foreshortened-cilia phenotype of dyf-6 mutants.
Reason: Core biological process; supported by the dyf-6 mutant cilium phenotype (PMID:16648645).
Supporting Evidence:
PMID:16648645
the cilia of the amphid and phasmid dendritic endings are foreshortened
GO:0043204 perikaryon
EXP
PMID:16648645
The molecular identities of the Caenorhabditis elegans intra...
KEEP AS NON CORE
Summary: Experimental localization of DYF-6 to the neuronal perikaryon (cell body) in addition to the cilium and dendrite. DYF-6::GFP is expressed throughout the ciliated sensory neurons.
Reason: Real experimental localization but not DYF-6's core functional site; the cell body reflects the site of expression rather than the IFT role.
Supporting Evidence:
PMID:16648645
DYF-6::GFP is expressed in amphid and phasmid neurons
GO:0030425 dendrite
IDA
PMID:16648645
The molecular identities of the Caenorhabditis elegans intra...
KEEP AS NON CORE
Summary: WormBase IDA localization of DYF-6 to the sensory dendrites, the route along which IFT cargo travels between the cell body and the ciliated ending.
Reason: Real localization along the trafficking route to the cilium, but not the core functional site (the cilium/IFT machinery).
GO:0042073 intraciliary transport
IDA
PMID:16648645
The molecular identities of the Caenorhabditis elegans intra...
ACCEPT
Summary: Direct observation that DYF-6::GFP moves within the ciliated endings of amphid and phasmid neurons demonstrates that DYF-6 participates in intraflagellar (intraciliary) transport. This is the primary experimental basis for the core function.
Reason: Core biological process; strongest, directly observed evidence (PMID:16648645).
Supporting Evidence:
PMID:16648645
Movement of DYF-6::GFP within the ciliated endings of the neurons indicates that DYF-6 is involved in IFT
GO:0043025 neuronal cell body
IDA
PMID:16648645
The molecular identities of the Caenorhabditis elegans intra...
KEEP AS NON CORE
Summary: WormBase IDA localization of DYF-6 to the neuronal cell body, consistent with the EXP perikaryon annotation and with expression throughout ciliated sensory neurons.
Reason: Real localization but not the core functional site; reflects the neuron of expression rather than the IFT role.
Supporting Evidence:
PMID:16648645
DYF-6::GFP is expressed in amphid and phasmid neurons
GO:1905515 non-motile cilium assembly
IMP
PMID:16648645
The molecular identities of the Caenorhabditis elegans intra...
ACCEPT
Summary: C. elegans sensory cilia are non-motile, and DYF-6 is required to build them: dyf-6 mutants have foreshortened amphid and phasmid ciliary endings. This is the most specific and organism-appropriate biological-process term for DYF-6.
Reason: Core biological process; the non-motile-cilium wording matches the worm sensory cilium and is directly supported by the mutant phenotype (PMID:16648645).
Supporting Evidence:
PMID:16648645
the cilia of the amphid and phasmid dendritic endings are foreshortened

Core Functions

DYF-6/IFT46 is a structural subunit of intraflagellar transport (IFT) complex B in ciliated sensory neurons. It localizes to the cilium and the ciliary base (basal body / transition-zone region), undergoes bidirectional IFT movement within the ciliated dendritic endings, and is required to build full-length non-motile sensory cilia β€” dyf-6 loss foreshortens the amphid and phasmid ciliary endings and produces the OSM-6 mislocalization pattern typical of IFT complex B mutants. It has no catalytic domain and acts as a structural/adaptor component of IFT-B; no informative molecular-function term is currently assignable (MF-dark; see knowledge_gaps).

Supporting Evidence:
  • PMID:16648645
    Movement of DYF-6::GFP within the ciliated endings of the neurons indicates that DYF-6 is involved in IFT
  • PMID:16648645
    the cilia of the amphid and phasmid dendritic endings are foreshortened
  • PMID:28479320
    intraflagellar transport (IFT)-B complex abolishes dynein-2's ciliary localization

References

Gene Ontology annotation through association of InterPro records with GO terms
Annotation inferences using phylogenetic trees
Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location vocabulary mapping, accompanied by conservative changes to GO terms applied by UniProt
Electronic Gene Ontology annotations created by ARBA machine learning models
The molecular identities of the Caenorhabditis elegans intraflagellar transport genes dyf-6, daf-10 and osm-1.
Dynein-Driven Retrograde Intraflagellar Transport Is Triphasic in C. elegans Sensory Cilia.
Mutations affecting the chemosensory neurons of Caenorhabditis elegans.
file:worm/dyf-6/dyf-6-deep-research-falcon.md
Deep research report (Falcon/Edison): DYF-6/IFT46 in Caenorhabditis elegans

Suggested Questions for Experts

Q: Does C. elegans DYF-6/IFT46 act as a cargo adaptor within IFT-B (as Chlamydomonas IFT46 does for outer dynein arms), or purely as a structural scaffold, and which IFT-B subunits does it directly contact?

Suggested Experiments

Experiment: Affinity purification / cross-linking mass spectrometry and structural modeling of DYF-6 within the worm IFT-B complex to define its direct binding partners and position in the particle.

Type: interaction mapping

Experiment: Isoform-specific rescue and separation-of-function alleles (including N-terminal deletions) to test for a cargo-adaptor role and to determine whether the long dyf-6/F46F6.3 fusion isoform has any distinct function.

Type: genetics

Knowledge Gaps

What is not known β€” curated, literature-grounded statements of the open unknowns (the inverse of core functions).

Gap: The molecular function of DYF-6/IFT46 is undefined. It has a large disordered N-terminus and no recognizable catalytic domain, and no GO molecular-function term is assigned to it in C. elegans. Whether it acts as a cargo adaptor (Chlamydomonas IFT46 uses its N-terminal region to load outer dynein arms) or purely as an IFT-B scaffold, and which IFT-B subunit(s) it directly contacts in the worm, is not established.

OPEN ONTOLOGYBIOLOGY MF_DARK

What is known: DYF-6 is firmly established as an IFT complex B subunit that localizes to the cilium and ciliary base, undergoes IFT movement, and is required to build full-length non-motile sensory cilia. In other organisms the conserved C-terminal IFT46_B_C domain mediates direct binding to IFT52 (and, via a ternary module, IFT88) to build the IFT-B1 core, while the N-terminal region binds the ODA16 cargo adaptor to load outer dynein arms β€” a role restricted to motile cilia and therefore not expected in the worm. What is missing is a GO molecular-function representation of this structural/scaffolding activity (the MF aspect cannot express "structural constituent of IFT particle B"), and the specific IFT-B contacts and any cargo-adaptor role of DYF-6 have not been demonstrated for the C. elegans protein.

Significance: IFT46 is a conserved core IFT-B protein; defining its molecular activity (scaffolding vs a specific cargo-adaptor interaction) is central to understanding IFT-B architecture and cargo selection in cilia.

What would resolve it: Map DYF-6's direct IFT-B interaction partners by biochemistry/structure; test for cargo-adaptor activity; introduce a molecular-function term for an IFT-particle structural/adaptor subunit activity.

Provenance (the field's own admissions):

Proposed term (ontology gap):

Gap: dyf-6 is a complex locus that, besides the short conserved IFT46 isoforms, produces a longer nematode-specific transcript fusing the dyf-6 ORF to the upstream gene F46F6.3; the biological role of this long product is uncharacterized. All functional and rescue data derive from the short isoforms, so whether the long isoform contributes anything beyond the conserved IFT-B function is unknown.

OPEN BIOLOGY RESIDUAL_SUBGAP

What is known: The short (conserved) dyf-6 isoforms confer full dye-filling rescue and undergo IFT; the F46F6.3-fused long transcript is nematode-specific and has no assigned function, and existing alleles/reporters primarily report on the short forms.

Significance: Establishing whether the nematode-specific long isoform has a distinct role would clarify whether the dyf-6 locus encodes more than the conserved IFT-B subunit function.

What would resolve it: Isoform-specific knockouts/rescue and expression analysis of the long dyf-6/F46F6.3 fusion product.

Provenance (the field's own admissions):

Deep Research

Falcon

(dyf-6-deep-research-falcon.md)
Comprehensive Research Report: DYF-6/IFT46 in *Caenorhabditis elegans* Falcon Edison Scientific Literature 40 citations 2 artifacts 2026-07-04T19:21:23.496379

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.

Comprehensive Research Report: DYF-6/IFT46 in Caenorhabditis elegans

1. Gene Identity and Protein Overview

The dyf-6 (abnormal dye filling protein 6) gene in Caenorhabditis elegans (ORF: F46F6.4; UniProt: Q0G838) encodes the worm ortholog of Intraflagellar Transport Protein 46 (IFT46). The protein belongs to the IFT46 family and contains a conserved Intraflagellar Transport Complex B C-terminal domain (IFT46_B_C; PF12317/IPR022088). DYF-6/IFT46 is a structural subunit of the intraflagellar transport (IFT) machinery, functioning as a core component of the IFT-B complex that is essential for the assembly, maintenance, and cargo transport functions of cilia in sensory neurons.

2. Primary Function: Structural Adapter in Intraflagellar Transport

DYF-6/IFT46 does not function as an enzyme, transporter, or signaling molecule in the conventional sense. Rather, it serves as a structural and adapter subunit of the IFT-B complex, an essential multi-protein machine that mediates bidirectional transport of cargo along ciliary axonemal microtubules. Cilia lack protein synthesis machinery, and thus all ciliary proteins must be synthesized in the cell body and transported into the cilium by IFT (lv2017intraflagellartransportprotein pages 4-7). DYF-6/IFT46 fulfills dual roles within this system:

2.1 Core IFT-B Complex Assembly

IFT46 is a pivotal subunit of the IFT-B1 core subcomplex, which consists of approximately 10 subunits: IFT22, IFT25, IFT27, IFT46, IFT52, IFT56, IFT70, IFT74, IFT81, and IFT88 (liu2025structuremakesa pages 1-2, nakayama2018ciliaryproteintrafficking pages 3-3). Within this core, IFT46 belongs to the B1-2 subgroup alongside IFT52, IFT56, IFT70, and IFT88 (nakayama2018ciliaryproteintrafficking pages 3-3). IFT46 directly interacts with IFT52 through large hydrophobic surfaces at their carboxy-terminal domains, and these two proteins together interact with IFT88 to form a stable ternary complex (IFT46–IFT52–IFT88) that constitutes a critical core module of IFT-B (lucker2010directinteractionsof pages 9-9, taschner2016theintraflagellartransport pages 5-6). IFT46 plays a stabilization role for both IFT52 and IFT88 within this complex (lucker2010directinteractionsof pages 9-9).

2.2 Cargo Adapter for Outer Dynein Arm Transport

In organisms with motile cilia, IFT46 has a specialized function in transporting outer dynein arms (ODAs) via its interaction with the cargo adapter ODA16 (DAW1 in mammals). The N-terminal domain of IFT46 (approximately amino acids 1–147 in Chlamydomonas), which is predicted to be intrinsically disordered, binds directly to ODA16. ODA16 is a WD-repeat protein whose eight-bladed Ξ²-propeller contains a structural cleft that accommodates IFT46's unstructured N-terminal domain (lechtreck2022cargoadaptersexpand pages 3-4, lechtreck2022cargoadaptersexpand pages 2-3). This ODA16–IFT46 interaction is essential for efficient ODA transport to the ciliary tip. In Chlamydomonas, mutants expressing N-terminally truncated IFT46 can assemble flagella but these flagella specifically lack most outer dynein arms (nakayama2018ciliaryproteintrafficking pages 4-5, fassad2017c11orf70mutationscausing pages 40-46). Notably, the direct ODA16–IFT46 interaction has not been demonstrated in human cells, where additional factors may be required (huang2023arl3regulatesoda16mediated pages 8-12).

Since C. elegans possesses only non-motile sensory cilia that lack dynein arms, the ODA transport function of IFT46's N-terminus may be less relevant in nematodes, though the protein's core IFT-B assembly function is fully conserved.

3. Subcellular Localization

DYF-6/IFT46 localizes to two principal subcellular compartments: the basal body (the ciliary base) and the cilium itself. Detailed studies in Chlamydomonas demonstrated that YFP-tagged IFT46 concentrates at the basal body and shows punctate distribution along the length of the flagellum, consistent with its association with moving IFT trains (lv2017intraflagellartransportprotein pages 4-7, lucker2010directinteractionsof pages 6-6). IFT-B proteins, including IFT46, form a semi-circular tri-lobed arc at the basal body (lv2017intraflagellartransportprotein pages 4-7). Anterograde IFT trains containing IFT-B proteins measure approximately 233 nm and move along B-microtubules of the axoneme (lv2017intraflagellartransportprotein pages 4-7).

The basal body localization of IFT46 depends critically on IFT52 but not vice versa, establishing a hierarchical recruitment mechanism. IFT52 and IFT46 preassemble as subcomplexes in the cytoplasm or at the trans-Golgi network (TGN) before being delivered to the basal body through vesicular or non-vesicle-mediated transport pathways (lv2017intraflagellartransportprotein pages 40-45, lv2017intraflagellartransportprotein pages 11-14). The C-terminal sequence of IFT46 (amino acids 246–321, designated BBTS3) serves as the basal body targeting sequence, which is also necessary for ciliary targeting (lv2017intraflagellartransportprotein pages 1-4). The specific leucine residues L285 and L286 within IFT46 are critical for the binding interface with IFT52 and, consequently, for proper basal body localization (lv2017intraflagellartransportprotein pages 9-11, lv2017intraflagellartransportprotein pages 11-14).

In C. elegans, DYF-6 functions in the sensory cilia of amphid and phasmid neurons, where it participates in IFT-mediated protein trafficking along the axoneme (cevik2013activetransportand pages 6-8, cevik2013activetransportand pages 8-10).

4. Protein Interactions and Pathway Involvement

The known protein-protein interactions of DYF-6/IFT46 are summarized below:

Interaction Partner Binding Domain on IFT46 Organism Studied Functional Role of Interaction Key Reference
IFT52 C-terminal domain of IFT46; hydrophobic interface, including residues around L285/L286 important for recruitment Chlamydomonas reinhardtii; human/cross-species IFT studies Direct IFT46-IFT52 binding supports IFT-B1 core complex assembly and recruits IFT46 to the basal body/ciliary base before train assembly (lv2017intraflagellartransportprotein pages 35-40, lv2017intraflagellartransportprotein pages 40-45, lv2017intraflagellartransportprotein pages 9-11, lv2017intraflagellartransportprotein pages 11-14, lv2017intraflagellartransportprotein pages 1-4) Lv et al., 2017; Taschner & Lorentzen, 2016 (lv2017intraflagellartransportprotein pages 35-40, lv2017intraflagellartransportprotein pages 1-4, taschner2016theintraflagellartransport pages 5-6)
IFT88 No independent IFT46-only binding domain resolved here; interacts as part of an IFT46-IFT52-IFT88 ternary core complex Chlamydomonas reinhardtii Stabilizes the IFT-B core architecture; IFT46, IFT52, and IFT88 form a direct ternary complex essential for core IFT-B complex integrity (lucker2010directinteractionsof pages 9-9, lucker2010directinteractionsof pages 1-1) Lucker et al., 2010 (lucker2010directinteractionsof pages 9-9)
ODA16/DAW1 N-terminal domain of IFT46, especially aa 1-147 in Chlamydomonas; not clearly conserved in human DAW1-IFT46 binding Chlamydomonas reinhardtii; comparative human studies Cargo-adapter interaction for outer dynein arm (ODA) transport into cilia/flagella; truncation of the IFT46 N-terminus impairs ODA transport and causes axonemes lacking most ODAs (lechtreck2022cargoadaptersexpand pages 3-4, lechtreck2022cargoadaptersexpand pages 2-3, huang2023arl3regulatesoda16mediated pages 8-12, nakayama2018ciliaryproteintrafficking pages 4-5, fassad2017c11orf70mutationscausing pages 40-46) Lechtreck, 2022; Wang et al., 2020; Nakayama & Katoh, 2018 (lechtreck2022cargoadaptersexpand pages 3-4, huang2023arl3regulatesoda16mediated pages 8-12, nakayama2018ciliaryproteintrafficking pages 4-5)
ARL13B Indirectly via the IFT46-IFT56 dimer/subcomplex rather than a mapped standalone IFT46 motif human; Caenorhabditis elegans Supports ciliary membrane protein localization/retention and ciliary trafficking regulation; ARL13B/ARL-13 associates with IFT-B through IFT46-IFT56, and dyf-6/IFT46 affects ARL-13 compartmentalization in worm cilia (nozaki2017regulationofciliary pages 4-7, nozaki2017regulationofciliary pages 31-35, cevik2013activetransportand pages 10-11, cevik2013activetransportand pages 8-10, cevik2013activetransportand pages 6-8) Cevik et al., 2013; Nozaki et al., 2017 (nozaki2017regulationofciliary pages 4-7, cevik2013activetransportand pages 10-11, nozaki2017regulationofciliary pages 7-10)
IFT81/IFT74 Via higher-order IFT-B core interactions; IFT46 pairs with IFT52, which associates with the IFT81/IFT74 module during core assembly Multiple organisms Promotes IFT-B core complex formation; IFT46-IFT52 associates with IFT81/IFT74 to build the core scaffold that underlies anterograde IFT train assembly (lv2017intraflagellartransportprotein pages 9-11, liu2025structuremakesa pages 1-2, nakayama2018ciliaryproteintrafficking pages 3-3, taschner2016theintraflagellartransport pages 5-6, lucker2010directinteractionsof pages 9-9) Taschner & Lorentzen, 2016; Nakayama & Katoh, 2018; Lucker et al., 2010 (nakayama2018ciliaryproteintrafficking pages 3-3, taschner2016theintraflagellartransport pages 5-6, lucker2010directinteractionsof pages 9-9)

Table: This table summarizes the main known DYF-6/IFT46 interaction partners, the mapped or inferred IFT46 binding regions, and the functional significance of each interaction in IFT-B assembly, cargo transport, and ciliary localization.

4.1 IFT46–IFT56 Dimer and ARL13B Regulation

A particularly important interaction in the context of C. elegans sensory cilia is the association of IFT46 with IFT56 to form a heterodimer that serves as the binding site for ARL13B (ARL-13 in C. elegans), a Joubert syndrome-associated small GTPase critical for ciliary membrane composition (nozaki2017regulationofciliary pages 4-7, nozaki2017regulationofciliary pages 31-35, nozaki2017regulationofciliary pages 7-10). In C. elegans, dyf-6 mutants show reduced ARL-13 at ciliary membranes and mislocalization of ARL-13 to the periciliary membrane, indicating that DYF-6/IFT46 is essential for maintaining ARL-13 within its proper ciliary membrane subdomain through active transport mechanisms (cevik2013activetransportand pages 6-8). FRAP analyses revealed that in dyf-6 mutants, ARL-13 shows slow diffusion rates between ciliary and periciliary membrane compartments, suggesting intact transition zone barriers but defective active transport (cevik2013activetransportand pages 6-8, cevik2013activetransportand pages 8-10).

5. Functional Domains of DYF-6/IFT46

The distinct functional regions of DYF-6/IFT46 highlight how the protein's N-terminal and C-terminal halves serve separable functions in cargo transport and complex assembly, respectively:

Domain/Region Amino Acid Range (approximate) Function Evidence
N-terminal domain aa 1-147 (Chlamydomonas IFT46) Predicted intrinsically disordered region that binds the cargo adapter ODA16 and is required for efficient outer dynein arm (ODA) transport; this N-terminal ODA16-binding function is conserved mainly in organisms with motile cilia Reviews and primary studies describe direct ODA16 interaction with the IFT46 N-terminus, and truncation causes strong ODA loss from flagella (lechtreck2022cargoadaptersexpand pages 3-4, lechtreck2022cargoadaptersexpand pages 2-3, nakayama2018ciliaryproteintrafficking pages 4-5, fassad2017c11orf70mutationscausing pages 40-46)
Assembly-critical internal segment aa 26-50 Required for functional rescue of flagellar assembly in Chlamydomonas ift46 mutants; indicates this short N-proximal segment contributes to core ciliogenic activity beyond the extreme N-terminus Recombinant rescue experiments showed aa 26-50 are necessary for flagellar assembly rescue, whereas the first 25 aa are dispensable (lucker2010directinteractionsof pages 9-9)
C-terminal domain / BBTS3 aa 246-321 Basal body targeting sequence (BBTS3); mediates recruitment to basal bodies/ciliary base, supports interaction with IFT52, and corresponds to the conserved IFT46_B_C region important for IFT-B incorporation and trafficking Localization and mutational studies mapped the basal body targeting sequence to the C-terminus and showed direct functional coupling to IFT52 (lv2017intraflagellartransportprotein pages 9-11, lv2017intraflagellartransportprotein pages 11-14, lv2017intraflagellartransportprotein pages 4-7, lv2017intraflagellartransportprotein pages 1-4)
IFT52-binding interface residues L285/L286 Critical residues within the C-terminal region for IFT52 binding; disruption impairs IFT46 recruitment/localization and the IFT52-IFT46 interaction Point-mutation analysis identified L285/L286 as essential for the IFT52 interaction interface (lv2017intraflagellartransportprotein pages 40-45, lv2017intraflagellartransportprotein pages 9-11, lv2017intraflagellartransportprotein pages 11-14, lv2017intraflagellartransportprotein pages 1-4)
Full C-terminus Broadly the C-terminal half, including the terminal interaction surface Required for IFT-B complex stability and assembly through hydrophobic interactions with IFT52; supports formation of the IFT46-IFT52-IFT88 core module and incorporation into the IFT-B1 complex Structural/biochemical studies show the C-terminus stabilizes IFT-B core assembly via IFT52 interaction and contributes to ternary complex formation with IFT88 (lucker2010directinteractionsof pages 9-9, taschner2016theintraflagellartransport pages 5-6)

Table: This table summarizes the main functional regions of IFT46/DYF-6, linking specific sequence segments to basal body targeting, IFT-B complex assembly, and cargo-adapter interactions. It is useful for distinguishing the conserved ciliogenic core functions of the protein from the motile-cilia-specific ODA transport role of its N-terminus.

6. Mutant Phenotypes and Experimental Evidence

6.1 Chlamydomonas IFT46 Mutants

The ift46-1 null mutant in Chlamydomonas reinhardtii produces stunted, paralyzed flagella significantly shorter than wild-type cells, with daughter cells remaining restricted within the mother cell wall (lv2017intraflagellartransportprotein pages 4-7). The ift46-2 strain carries a deletion of most of the IFT46 gene and displays a predominantly bald (non-flagellated) phenotype, with only approximately 6% of cells assembling short flagella averaging 3 Β΅m in length (lucker2010directinteractionsof pages 1-1, lucker2010directinteractionsof pages 6-6, lucker2010directinteractionsof pages 5-6). This phenotype is slightly less severe than ift52 or ift88 mutants (lucker2010directinteractionsof pages 1-1). Electroporation of recombinant IFT46 protein successfully rescued the flagellar assembly defect, restoring motile flagella within 4 hours post-electroporation, including normal photophobic and phototactic responses (lucker2010directinteractionsof pages 6-6, lucker2010directinteractionsof pages 5-6). Rescue experiments further demonstrated that amino acids 26–50 are required for flagellar assembly function, whereas the first 25 amino acids are dispensable (lucker2010directinteractionsof pages 9-9).

6.2 C. elegans DYF-6 Mutant Phenotypes

In C. elegans, dyf-6 mutants exhibit the characteristic dye-filling defective (Dyf) phenotype, which is the hallmark of ciliary dysfunction in worm sensory neurons. The Dyf phenotype indicates that the amphid and phasmid cilia are structurally compromised, preventing uptake of lipophilic fluorescent dyes such as DiI that normally enter neurons through intact sensory cilia (cevik2013activetransportand pages 8-10, cevik2013activetransportand pages 6-8). The study by Cevik et al. (2013) demonstrated that in dyf-6 IFT-B mutants, the Joubert syndrome protein ARL-13 is mislocalized, with reduced amounts in the ciliary middle segment and aberrant accumulation at periciliary membranes (cevik2013activetransportand pages 6-8). Additionally, dyf-6 mutants display defects in IFT-dependent anthelmintic drug uptake through amphid sensory cilia, as IFT genes including dyf-6 are among the ciliary genes required for proper avermectin sensitivity in C. elegans (brinzer2021theuptakeof pages 10-12, brinzer2021theuptakeof pages 14-17).

7. Transcriptional Regulation

Expression of ciliary genes in C. elegans, including those encoding IFT-B complex components, is regulated by the RFX transcription factor DAF-19, which binds to conserved X-box motifs in promoter regions. DAF-19 is essential for ciliary gene expressionβ€”when DAF-19 is non-functional, ciliated neurons lose their cilia and exhibit sensory defects (chu2012finetuningof pages 1-2). C. elegans possesses approximately 60 ciliated sensory neurons organized in three main clusters: labial and amphid neurons in the head, and phasmid neurons in the tail (warrington2018computationalandmolecular pages 21-25). The dyf-6 gene, like other ciliary genes, is expected to be under DAF-19/RFX transcriptional control given that its homolog CG15161 in Drosophila has been identified as an RFX target gene (chu2012finetuningof pages 1-2). Multiple X-box motifs can cooperate to fine-tune the expression levels of ciliary genes in specific ciliated neuron subtypes (chu2012finetuningof pages 5-8).

8. Evolutionary Conservation

IFT46 is broadly conserved across eukaryotes that possess cilia, from the green alga Chlamydomonas reinhardtii to nematodes, zebrafish, and mammals. The C-terminal IFT46_B_C domain (PF12317) that mediates IFT-B complex interactions is conserved across all species, while the N-terminal region involved in ODA16 binding is specifically conserved in organisms with motile cilia (lechtreck2022cargoadaptersexpand pages 3-4, lechtreck2022cargoadaptersexpand pages 2-3). In zebrafish, IFT46 plays an essential role in cilia development, and mutations lead to typical ciliopathy-related phenotypes. In human cells, IFT46 is part of the IFT-B1 core subcomplex where it participates in the same core interactions with IFT52 and IFT88, though the ODA16-IFT46 interaction for dynein transport may involve additional factors compared to Chlamydomonas (huang2023arl3regulatesoda16mediated pages 8-12).

9. Summary

DYF-6/IFT46 in C. elegans is a core structural subunit of the IFT-B1 complex that is indispensable for intraflagellar transport and sensory cilium assembly. It localizes to the basal body and the ciliary axoneme of sensory neurons, where it functions as part of the IFT machinery that transports ciliary proteins between the cell body and the cilium. Its C-terminal domain mediates incorporation into the IFT-B core complex through direct interaction with IFT52, while its N-terminal domain serves as a binding site for the cargo adapter ODA16 in organisms with motile cilia. Through its interaction with IFT56, IFT46 also contributes to the localization and retention of ciliary membrane proteins such as ARL13B. Loss of DYF-6 in C. elegans results in dye-filling defects indicative of structurally compromised sensory cilia, mislocalization of ciliary membrane proteins, and consequent sensory deficits. The gene's expression is regulated by the ciliary transcription factor DAF-19/RFX, placing it within the core ciliogenesis transcriptional program.

References

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Artifacts

Citations

  1. lv2017intraflagellartransportprotein pages 4-7
  2. nakayama2018ciliaryproteintrafficking pages 3-3
  3. lucker2010directinteractionsof pages 9-9
  4. lv2017intraflagellartransportprotein pages 1-4
  5. cevik2013activetransportand pages 6-8
  6. lucker2010directinteractionsof pages 1-1
  7. chu2012finetuningof pages 1-2
  8. warrington2018computationalandmolecular pages 21-25
  9. chu2012finetuningof pages 5-8
  10. liu2025structuremakesa pages 1-2
  11. taschner2016theintraflagellartransport pages 5-6
  12. lechtreck2022cargoadaptersexpand pages 3-4
  13. lechtreck2022cargoadaptersexpand pages 2-3
  14. nakayama2018ciliaryproteintrafficking pages 4-5
  15. lucker2010directinteractionsof pages 6-6
  16. lv2017intraflagellartransportprotein pages 40-45
  17. lv2017intraflagellartransportprotein pages 11-14
  18. lv2017intraflagellartransportprotein pages 9-11
  19. cevik2013activetransportand pages 8-10
  20. lv2017intraflagellartransportprotein pages 35-40
  21. nozaki2017regulationofciliary pages 4-7
  22. nozaki2017regulationofciliary pages 31-35
  23. cevik2013activetransportand pages 10-11
  24. nozaki2017regulationofciliary pages 7-10
  25. lucker2010directinteractionsof pages 5-6
  26. brinzer2021theuptakeof pages 10-12
  27. brinzer2021theuptakeof pages 14-17
  28. https://doi.org/10.1242/jcs.200758,
  29. https://doi.org/10.1002/cm.70033,
  30. https://doi.org/10.1093/jb/mvx087,
  31. https://doi.org/10.1074/jbc.m110.106997,
  32. https://doi.org/10.1101/cshperspect.a028092,
  33. https://doi.org/10.1242/jcs.260408,
  34. https://doi.org/10.1101/211953,
  35. https://doi.org/10.1101/2023.04.12.536397,
  36. https://doi.org/10.1371/journal.pgen.1003977,
  37. https://doi.org/10.1242/jcs.197004,
  38. https://doi.org/10.1101/2021.10.22.465401,
  39. https://doi.org/10.1093/nar/gkr690,
  40. https://doi.org/10.48550/arxiv.1810.00478,

πŸ“š Additional Documentation

Notes

(dyf-6-notes.md)

dyf-6 (Q0G838) research notes

Identity (from UniProt Q0G838, IFT46_CAEEL)

  • Gene: dyf-6 (F46F6.4), WormBase WBGene00001122, chromosome X. "dyf" = abnormal dye filling.
  • Protein: Intraflagellar transport protein 46 homolog (IFT46). 471 aa (isoform a). Human ortholog IFT46 = Q9NQC8; mouse Q9CQ63 (used in IBA WITH/FROM).
  • Domain/family: PANTHER PTHR13376 (INTRAFLAGELLAR TRANSPORT PROTEIN 46 HOMOLOG); Pfam PF12317 (IFT46_B_C); InterPro IPR022088 (Intraflagellar_transp_cmplxB). Large N-terminal disordered/acidic region (res 1–202); no catalytic domain. So DYF-6 is MF-dark: a structural/adaptor subunit of IFT-B with no assigned molecular activity (unlike its worm paralog-family neighbor dyf-11/IFT54, which carries an InterPro2GO microtubule-binding transfer β€” dyf-6 has NO MF annotation at all in GOA).
  • Complex: Component of IFT complex B (ComplexPortal CPX-1290). UniProt SUBUNIT (from PMID:28479320): "Component of the IFT complex B composed of at least che-2, che-13, dyf-1, dyf-3, dyf-6, dyf-11, dyf-13, ift-20, ift-74, ift-81, ifta-2, osm-1, osm-5 and osm-6."
  • Localization (UniProt, from PMID:16648645): Cell projection, cilium; Cytoplasm/cytoskeleton, cilium basal body; Cell projection, dendrite; Perikaryon. "Highly expressed in the transition zones between the cilium basal body and the dendrites."
  • Isoforms: 4 (a Q0G838-1 displayed; b/c/d). dyf-6 is a complex locus; a longer mRNA fuses the dyf-6 ORF to the upstream gene F46F6.3 β€” the short forms confer full rescue and are the conserved/functional forms.

KNOWN (well-supported)

  1. DYF-6 is an IFT (intraflagellar transport) protein that moves within cilia.
  2. PMID:16648645
  3. PMID:16648645

  4. DYF-6 is (very likely) an IFT complex B component. Bell 2006 inferred this from the OSM-6::GFP mislocalization pattern; Yi 2017/ComplexPortal biochemically place it in IFT-B.

  5. PMID:16648645
  6. PMID:16648645
  7. UniProt SUBUNIT + ComplexPortal CPX-1290 (from PMID:28479320) list dyf-6 in IFT-B.

  8. DYF-6 is required to build/maintain sensory cilia; loss foreshortens amphid & phasmid cilia.

  9. PMID:16648645
  10. C. elegans sensory cilia are non-motile, so the specific process term is non-motile cilium assembly (GO:1905515, WormBase IMP).

  11. DYF-6 acts cell-autonomously in the amphid sensory neurons; loss causes dye-filling/chemotaxis defects.

  12. PMID:16648645
  13. PMID:16648645

  14. Expression: DYF-6::GFP in amphid & phasmid (and IL-region) ciliated neurons, plus hypodermis; expressed hatching→adult incl. dauer.

  15. PMID:16648645
  16. PMID:16648645

  17. Conservation: orthologs in fly (CG15161, ciliary) and mammals; human ortholog conserved throughout.

  18. PMID:16648645
  19. PMID:16648645

  20. DYF-6/IFT-B is needed for dynein-2 ciliary entry (retrograde IFT context).

  21. PMID:28479320 (abstract-only cache; full text is the source of the IFT-B mass-spec membership)

NOT known / gaps

  • Molecular function of DYF-6/IFT46 is undefined. No catalytic domain; large disordered N-terminus. Which IFT-B subunit(s) it directly contacts in the worm, and whether it (like Chlamydomonas IFT46, which uses its N-terminal domain to load outer dynein arms) has a cargo-adaptor activity, is not established here. GO MF aspect cannot currently express "structural subunit of IFT-B" (ontology gap). PMID:16648645
  • Function of the long dyf-6/F46F6.3 fusion isoform is unknown. PMID:16648645
  • Whether DYF-6 has any non-ciliary role (it is also seen in dendrite/perikaryon/cell body) is untested.

Annotation-review plan (19 GOA annotations)

  • CORE (ACCEPT): intraciliary transport particle B (IBA part_of; NAS part_of), intraciliary transport (IBA/IEA/NAS/IDA involved_in), cilium assembly (IBA/NAS involved_in), non-motile cilium assembly (IMP involved_in), cilium (IEA/NAS located_in).
  • MODIFY: microtubule organizing center (IBA is_active_in) β†’ ciliary basal body GO:0036064 (documented basal-body/transition-zone localization).
  • MARK_AS_OVER_ANNOTATED: motile cilium (IBA is_active_in) β€” C. elegans has NO motile cilia (mammalian-biased IBA over-propagation; worm cilia are non-motile sensory); plasma membrane bounded cell projection GO:0120025 (ARBA IEA) β€” over-general grouping parent of cilium+dendrite already annotated.
  • KEEP_AS_NON_CORE (real but not core location/route): dendrite (IEA/IDA), perikaryon (IEA/EXP), neuronal cell body (IDA).

Provenance note

Cache: PMID:16648645 full text available (Bell et al. 2006, Genetics, PMC1526656). PMID:28479320 abstract-only (Yi et al. 2017, Curr Biol) β€” IFT-B membership rests on full-text mass-spec not in cache; annotations from it are ComplexPortal NAS, accepted deferring to curator/ComplexPortal.

Falcon deep research (dyf-6-deep-research-falcon.md, 2026-07-04) β€” additional context

Genuine, gene-correct report (40 citations, 2 artifacts). Key points that refine but do not
contradict the review:

  • IFT-B1 core architecture: IFT46's conserved C-terminal IFT46_B_C domain (PF12317) directly
    binds IFT52, and IFT46–IFT52–IFT88 form a stable ternary core module of IFT-B1 (Lucker et al.
    2010 J Biol Chem; Lv et al. 2017 J Cell Sci; Taschner & Lorentzen 2016). So DYF-6 is not
    partner-less β€” but these interactions are characterized in Chlamydomonas/human, not the worm,
    and GO MF has no term for the structural role. [falcon: "IFT46 directly interacts with IFT52
    through large hydrophobic surfaces at their carboxy-terminal domains"]
  • Basal-body targeting: IFT52 recruits IFT46 to the basal body; the IFT46 C-terminal BBTS3
    segment (~aa 246–321) is the basal-body/ciliary targeting sequence; L285/L286 are key for IFT52
    binding (Lv et al. 2017). Supports the MTOC→ciliary basal body MODIFY.
  • N-terminal ODA16 cargo-adaptor role is MOTILE-cilia-specific (Chlamydomonas ODA transport),
    hence NOT expected in C. elegans non-motile sensory cilia β€” independent support for marking the
    IBA "motile cilium" annotation as over-annotated. [falcon: "the N-terminal region involved in
    ODA16 binding is specifically conserved in organisms with motile cilia"; "Since C. elegans
    possesses only non-motile sensory cilia that lack dynein arms, the ODA transport function of
    IFT46's N-terminus may be less relevant in nematodes"]
  • Worm-specific ARL-13 trafficking (NOT in GOA): Cevik et al. 2013 report that dyf-6 mutants
    mislocalize the Joubert-syndrome protein ARL-13/ARL13B (reduced in ciliary middle segment,
    accumulates at periciliary membrane), via the IFT46–IFT56 dimer that binds ARL13B (Nozaki et al.
    2017). This is a genuine dyf-6 functional finding but the primary papers are not in the
    publications cache, so it is recorded here as context only (not added as verbatim-supported
    annotation).
  • Transcriptional control: ciliary genes incl. IFT-B are DAF-19/RFX (X-box) targets; dyf-6's
    Drosophila ortholog CG15161 is an RFX target (consistent with X-box regulation of ciliary genes).
  • Anthelmintic uptake: dyf-6 among ciliary genes needed for avermectin uptake via amphid cilia
    (Brinzer et al. 2021) β€” a downstream/pleiotropic sensory readout, not a core function.

Not independently re-verified (LLM synthesis with internal citation keys); used as supporting
context. The two ontology/biology knowledge gaps in the review reflect this: partners known in
other species but (a) no GO MF term and (b) worm-specific contacts/adaptor role unproven.

πŸ“„ View Raw YAML

id: Q0G838
gene_symbol: dyf-6
product_type: PROTEIN
status: COMPLETE
aliases:
- IFT46
- F46F6.4
taxon:
  id: NCBITaxon:6239
  label: Caenorhabditis elegans
description: >-
  dyf-6 encodes the Caenorhabditis elegans ortholog of intraflagellar transport
  protein 46 (IFT46), a structural subunit of intraflagellar transport (IFT)
  complex B. IFT-B, with IFT-A and the kinesin-2 and dynein-2 motors, drives the
  bidirectional movement of ciliary cargo that builds and maintains the cilia of
  ciliated sensory neurons. The 471-residue protein has a large N-terminal
  disordered/acidic region and no recognizable catalytic domain. DYF-6 is
  expressed in ciliated amphid and phasmid sensory neurons (and hypodermis),
  localizes to the cilium, the ciliary base/basal body region, dendrites and the
  neuronal cell body, and undergoes processive IFT movement within the ciliated
  dendritic endings. It is required to build full-length sensory cilia: in dyf-6
  mutants the amphid and phasmid ciliary endings are foreshortened, the IFT-B
  marker OSM-6 is mislocalized in a pattern typical of complex B mutants, and the
  animals are defective in dye filling and chemotaxis. The ciliary role is
  conserved, with orthologs in Drosophila (CG15161, expressed in sensory cilia)
  and mammals (human IFT46). C. elegans has only non-motile sensory cilia, so the
  cilium DYF-6 builds is a non-motile sensory cilium.
references:
- id: GO_REF:0000002
  title: Gene Ontology annotation through association of InterPro records with GO
    terms
  findings: []
  reference_review:
    relevance: MEDIUM
    correctness: VERIFIED
    review_notes: >-
      InterPro2GO mapping of the IFT complex B family (IPR022088) to intraciliary
      transport (GO:0042073). The family/domain identity (Pfam IFT46_B_C,
      PANTHER PTHR13376) is correct for DYF-6, and the transferred biological
      process is independently confirmed by C. elegans experimental data, so the
      transfer is sound.
- id: GO_REF:0000033
  title: Annotation inferences using phylogenetic trees
  findings: []
  reference_review:
    relevance: MEDIUM
    correctness: VERIFIED
    review_notes: >-
      Standard GO_Central IBA pipeline. The IFT-B membership, intraciliary
      transport and cilium-assembly transfers are all corroborated by worm
      experimental data (PMID:16648645). The "motile cilium" (GO:0031514)
      transfer is a mammalian-biased over-propagation: C. elegans has no motile
      cilia, only non-motile sensory cilia.
- id: GO_REF:0000044
  title: Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location
    vocabulary mapping, accompanied by conservative changes to GO terms applied by
    UniProt
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: >-
      Electronic mapping of UniProt subcellular-location keywords (cilium,
      dendrite, perikaryon) to GO CC terms. Redundant with the experimental
      WormBase/UniProt annotations from PMID:16648645 but not incorrect.
- id: GO_REF:0000117
  title: Electronic Gene Ontology annotations created by ARBA machine learning models
  findings: []
  reference_review:
    relevance: LOW
    correctness: LOW_QUALITY
    review_notes: >-
      ARBA machine-learning annotation to the over-general grouping term
      "plasma membrane bounded cell projection" (GO:0120025), whose informative
      descendants (cilium, dendrite) are already annotated. Uninformative for
      this protein.
- id: PMID:16648645
  title: The molecular identities of the Caenorhabditis elegans intraflagellar transport
    genes dyf-6, daf-10 and osm-1.
  findings: []
  reference_review:
    relevance: HIGH
    correctness: VERIFIED
    review_notes: >-
      Bell et al. 2006 (Genetics, PMC1526656), the primary cloning and functional
      characterization of dyf-6. Establishes DYF-6 as an IFT protein that moves
      within amphid/phasmid ciliated endings, that dyf-6 mutants have foreshortened
      cilia, that dyf-6 functions cell-autonomously in amphid sensilla, and that
      the protein has no recognized motifs but is conserved to flies and mammals.
      Source of the WormBase IDA/IMP/EXP annotations. (Cache exposes the abstract;
      the full text additionally infers IFT complex B membership from the OSM-6
      mislocalization pattern.)
- id: PMID:28479320
  title: Dynein-Driven Retrograde Intraflagellar Transport Is Triphasic in C. elegans
    Sensory Cilia.
  findings: []
  reference_review:
    relevance: MEDIUM
    correctness: VERIFIED
    review_notes: >-
      Yi et al. 2017 (Curr Biol). Source of the ComplexPortal (CPX-1290) NAS
      annotations placing DYF-6 in IFT complex B and of the UniProt SUBUNIT list.
      Cached record is abstract-only; the abstract confirms that an intact IFT-B
      complex is required for dynein-2 ciliary entry, and the full text is the
      basis (affinity purification / mass spectrometry) for DYF-6's IFT-B
      membership. NAS annotations accepted, deferring to ComplexPortal/curator.
- id: PMID:7705621
  title: Mutations affecting the chemosensory neurons of Caenorhabditis elegans.
  findings: []
  reference_review:
    relevance: MEDIUM
    correctness: VERIFIED
    review_notes: >-
      Starich et al. 1995 (Genetics), the genetic screen that isolated dyf-6 among
      the dye-filling-defective (Dyf) mutants and characterized its chemotaxis /
      dye-filling disruption phenotype; cited by UniProt as a FUNCTION and
      DISRUPTION PHENOTYPE reference for dyf-6. Cached record is abstract-only (the
      abstract summarizes the screen without naming dyf-6), so it is included as
      background establishing the loss-of-function phenotype rather than as a
      verbatim source for a specific annotation.
- id: file:worm/dyf-6/dyf-6-deep-research-falcon.md
  title: "Deep research report (Falcon/Edison): DYF-6/IFT46 in Caenorhabditis elegans"
  findings: []
  reference_review:
    relevance: MEDIUM
    correctness: UNVERIFIED
    review_notes: >-
      AI-generated (Falcon/Edison) deep-research synthesis. Corroborates the
      primary-literature-grounded review: DYF-6/IFT46 is a structural IFT-B1 core
      subunit whose conserved C-terminal IFT46_B_C domain mediates direct IFT52
      (and, via a ternary module, IFT88) binding, while the N-terminal ODA16
      cargo-adaptor role is confined to organisms with motile cilia and is not
      expected in C. elegans non-motile sensory cilia. Also surfaces worm-specific
      literature (Cevik et al. 2013 on ARL-13 ciliary membrane trafficking in
      dyf-6 mutants) not in the GOA set. Underlying claims are drawn from
      Chlamydomonas/human work cited by internal keys, not independently
      re-verified here; treated as supporting context, not primary evidence.
existing_annotations:
- term:
    id: GO:0005815
    label: microtubule organizing center
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: is_active_in
  review:
    summary: >-
      IFT46 orthologs concentrate at and inject IFT trains from the ciliary base.
      In C. elegans this is the basal body / transition-zone region; UniProt
      curates DYF-6 basal-body localization from PMID:16648645. The basal body is
      a type of microtubule organizing center, so the term is not wrong, but the
      specific "ciliary basal body" (GO:0036064) is the accurate, informative CC.
    action: MODIFY
    reason: >-
      Term too general. DYF-6's documented location at the ciliary base is better
      captured by ciliary basal body (GO:0036064) than by the generic
      microtubule organizing center.
    proposed_replacement_terms:
    - id: GO:0036064
      label: ciliary basal body
    propagation_review:
      root_cause: TERM_SCOPING_PROBLEM
      failure_modes:
      - GRANULARITY_MISMATCH
      source_entities:
      - source_id: PANTHER:PTN000332669
        source_label: IFT46 family node
        source_status: SUPPORTS_TRANSFER
        comment: >-
          The IFT46 family node correctly places DYF-6 at the microtubule
          organizing center (basal body), but the transferred term is coarser than
          the specific ciliary basal body location documented for the worm protein.
- term:
    id: GO:0030992
    label: intraciliary transport particle B
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: part_of
  review:
    summary: >-
      DYF-6/IFT46 is a subunit of intraflagellar transport complex B (IFT-B).
      This phylogenetic transfer is corroborated by Bell et al.'s inference from
      the OSM-6 mislocalization pattern and by biochemical assignment to
      ComplexPortal CPX-1290.
    action: ACCEPT
    reason: >-
      Core cellular-component identity of DYF-6 as an IFT-B structural subunit.
    supported_by:
    - reference_id: PMID:28479320
      supporting_text: intraflagellar transport (IFT)-B complex abolishes dynein-2's
        ciliary localization
      reference_section_type: ABSTRACT
    - reference_id: file:worm/dyf-6/dyf-6-deep-research-falcon.md
      supporting_text: IFT46 is a pivotal subunit of the IFT-B1 core subcomplex
      reference_section_type: OTHER
- term:
    id: GO:0060271
    label: cilium assembly
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: involved_in
  review:
    summary: >-
      IFT is required to assemble and maintain cilia; dyf-6 mutants have
      foreshortened amphid and phasmid ciliary endings, so DYF-6 is required for
      cilium assembly.
    action: ACCEPT
    reason: >-
      Core biological process, supported experimentally in worm (PMID:16648645).
      The more specific worm-appropriate term non-motile cilium assembly is also
      annotated (IMP).
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: the cilia of the amphid and phasmid dendritic endings are
        foreshortened
      reference_section_type: ABSTRACT
- term:
    id: GO:0031514
    label: motile cilium
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: is_active_in
  review:
    summary: >-
      This IBA transfer from mammalian IFT46 assigns a motile-cilium location.
      C. elegans, however, has no motile cilia β€” all its cilia, including the
      amphid and phasmid sensory cilia in which DYF-6 acts, are non-motile. The
      matching worm annotation is non-motile cilium assembly (GO:1905515, IMP).
    action: MARK_AS_OVER_ANNOTATED
    reason: >-
      Taxonomically inappropriate over-propagation: C. elegans lacks motile
      cilia. DYF-6 acts in non-motile sensory cilia; the generic ciliary location
      (cilium, GO:0005929) is separately and correctly annotated.
    propagation_review:
      root_cause: PROPAGATION_BAD
      failure_modes:
      - LINEAGE_OR_TAXON_MISMATCH
      source_entities:
      - source_id: PANTHER:PTN000332669
        source_label: IFT46 family node
        source_status: SUPPORTS_SOURCE_BUT_NOT_TARGET
        comment: >-
          The IFT46 family node includes vertebrate members that function in
          motile cilia, but the C. elegans ortholog acts only in non-motile
          sensory cilia, so the motile-cilium location does not transfer.
- term:
    id: GO:0042073
    label: intraciliary transport
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: involved_in
  review:
    summary: >-
      DYF-6 functions in intraflagellar (intraciliary) transport as an IFT-B
      subunit; DYF-6::GFP undergoes IFT movement within ciliated endings.
    action: ACCEPT
    reason: >-
      Core biological process, confirmed experimentally in worm (PMID:16648645).
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: Movement of DYF-6::GFP within the ciliated endings of the
        neurons indicates that DYF-6 is involved in IFT
      reference_section_type: ABSTRACT
- term:
    id: GO:0005929
    label: cilium
  evidence_type: IEA
  original_reference_id: GO_REF:0000044
  qualifier: located_in
  review:
    summary: >-
      Electronic mapping of the UniProt cilium subcellular-location keyword.
      DYF-6 is directly observed within cilia, where it undergoes IFT.
    action: ACCEPT
    reason: >-
      Correct core ciliary location, corroborated by experimental IFT movement
      within ciliated endings (PMID:16648645).
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: Movement of DYF-6::GFP within the ciliated endings of the
        neurons indicates that DYF-6 is involved in IFT
      reference_section_type: ABSTRACT
- term:
    id: GO:0030425
    label: dendrite
  evidence_type: IEA
  original_reference_id: GO_REF:0000044
  qualifier: located_in
  review:
    summary: >-
      DYF-6 is present in the sensory dendrites through which IFT cargo is
      trafficked to the ciliated ending; UniProt curates dendrite localization
      from PMID:16648645 and WormBase makes the same call by IDA.
    action: KEEP_AS_NON_CORE
    reason: >-
      Real localization (the dendritic route to the cilium) but not the core
      site of DYF-6 function, which is the cilium/IFT machinery.
- term:
    id: GO:0042073
    label: intraciliary transport
  evidence_type: IEA
  original_reference_id: GO_REF:0000002
  qualifier: involved_in
  review:
    summary: >-
      InterPro2GO transfer (IPR022088, IFT complex B) to intraciliary transport.
      Duplicates the IBA/NAS/IDA intraciliary-transport annotations and is
      correct.
    action: ACCEPT
    reason: >-
      Core biological process; redundant with experimental IDA (PMID:16648645)
      but consistent.
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: Movement of DYF-6::GFP within the ciliated endings of the
        neurons indicates that DYF-6 is involved in IFT
      reference_section_type: ABSTRACT
- term:
    id: GO:0043204
    label: perikaryon
  evidence_type: IEA
  original_reference_id: GO_REF:0000044
  qualifier: located_in
  review:
    summary: >-
      Electronic mapping of the UniProt perikaryon subcellular-location keyword,
      the same call made experimentally (EXP) from PMID:16648645. DYF-6 is present
      in the neuronal cell body/perikaryon in addition to the cilium.
    action: KEEP_AS_NON_CORE
    reason: >-
      Real localization but not the core functional site; the cell body reflects
      the neuron of expression rather than DYF-6's IFT role.
- term:
    id: GO:0120025
    label: plasma membrane bounded cell projection
  evidence_type: IEA
  original_reference_id: GO_REF:0000117
  qualifier: located_in
  review:
    summary: >-
      ARBA machine-learning annotation to a high-level grouping term that is the
      parent of both cilium and dendrite, which are already specifically
      annotated for DYF-6.
    action: MARK_AS_OVER_ANNOTATED
    reason: >-
      Over-general electronic grouping term; its informative descendants (cilium,
      dendrite) are already annotated, so it adds no information.
- term:
    id: GO:0005929
    label: cilium
  evidence_type: NAS
  original_reference_id: PMID:28479320
  qualifier: located_in
  review:
    summary: >-
      ComplexPortal (CPX-1290) assertion that DYF-6 localizes to the cilium,
      consistent with the experimental observation that DYF-6 undergoes IFT
      within cilia.
    action: ACCEPT
    reason: >-
      Correct core ciliary location.
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: Movement of DYF-6::GFP within the ciliated endings of the
        neurons indicates that DYF-6 is involved in IFT
      reference_section_type: ABSTRACT
- term:
    id: GO:0030992
    label: intraciliary transport particle B
  evidence_type: NAS
  original_reference_id: PMID:28479320
  qualifier: part_of
  review:
    summary: >-
      ComplexPortal (CPX-1290) assignment of DYF-6 to IFT complex B, based on the
      affinity-purification / mass-spectrometry of the worm IFT-B complex in Yi
      et al. 2017 and reflected in the UniProt SUBUNIT statement.
    action: ACCEPT
    reason: >-
      Core cellular-component identity; biochemically supported IFT-B membership.
    supported_by:
    - reference_id: PMID:28479320
      supporting_text: intraflagellar transport (IFT)-B complex abolishes dynein-2's
        ciliary localization
      reference_section_type: ABSTRACT
- term:
    id: GO:0042073
    label: intraciliary transport
  evidence_type: NAS
  original_reference_id: PMID:28479320
  qualifier: involved_in
  review:
    summary: >-
      ComplexPortal assertion that DYF-6, as an IFT-B subunit, functions in
      intraciliary transport, consistent with the worm experimental data.
    action: ACCEPT
    reason: >-
      Core biological process; consistent with IFT-B membership and experimental
      IFT movement (PMID:16648645).
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: Movement of DYF-6::GFP within the ciliated endings of the
        neurons indicates that DYF-6 is involved in IFT
      reference_section_type: ABSTRACT
- term:
    id: GO:0060271
    label: cilium assembly
  evidence_type: NAS
  original_reference_id: PMID:28479320
  qualifier: involved_in
  review:
    summary: >-
      ComplexPortal assertion that DYF-6 functions in cilium assembly, consistent
      with the foreshortened-cilia phenotype of dyf-6 mutants.
    action: ACCEPT
    reason: >-
      Core biological process; supported by the dyf-6 mutant cilium phenotype
      (PMID:16648645).
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: the cilia of the amphid and phasmid dendritic endings are
        foreshortened
      reference_section_type: ABSTRACT
- term:
    id: GO:0043204
    label: perikaryon
  evidence_type: EXP
  original_reference_id: PMID:16648645
  qualifier: located_in
  review:
    summary: >-
      Experimental localization of DYF-6 to the neuronal perikaryon (cell body)
      in addition to the cilium and dendrite. DYF-6::GFP is expressed throughout
      the ciliated sensory neurons.
    action: KEEP_AS_NON_CORE
    reason: >-
      Real experimental localization but not DYF-6's core functional site; the
      cell body reflects the site of expression rather than the IFT role.
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: DYF-6::GFP is expressed in amphid and phasmid neurons
      reference_section_type: ABSTRACT
- term:
    id: GO:0030425
    label: dendrite
  evidence_type: IDA
  original_reference_id: PMID:16648645
  qualifier: located_in
  review:
    summary: >-
      WormBase IDA localization of DYF-6 to the sensory dendrites, the route
      along which IFT cargo travels between the cell body and the ciliated
      ending.
    action: KEEP_AS_NON_CORE
    reason: >-
      Real localization along the trafficking route to the cilium, but not the
      core functional site (the cilium/IFT machinery).
- term:
    id: GO:0042073
    label: intraciliary transport
  evidence_type: IDA
  original_reference_id: PMID:16648645
  qualifier: involved_in
  review:
    summary: >-
      Direct observation that DYF-6::GFP moves within the ciliated endings of
      amphid and phasmid neurons demonstrates that DYF-6 participates in
      intraflagellar (intraciliary) transport. This is the primary experimental
      basis for the core function.
    action: ACCEPT
    reason: >-
      Core biological process; strongest, directly observed evidence
      (PMID:16648645).
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: Movement of DYF-6::GFP within the ciliated endings of the
        neurons indicates that DYF-6 is involved in IFT
      reference_section_type: ABSTRACT
- term:
    id: GO:0043025
    label: neuronal cell body
  evidence_type: IDA
  original_reference_id: PMID:16648645
  qualifier: located_in
  review:
    summary: >-
      WormBase IDA localization of DYF-6 to the neuronal cell body, consistent
      with the EXP perikaryon annotation and with expression throughout ciliated
      sensory neurons.
    action: KEEP_AS_NON_CORE
    reason: >-
      Real localization but not the core functional site; reflects the neuron of
      expression rather than the IFT role.
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: DYF-6::GFP is expressed in amphid and phasmid neurons
      reference_section_type: ABSTRACT
- term:
    id: GO:1905515
    label: non-motile cilium assembly
  evidence_type: IMP
  original_reference_id: PMID:16648645
  qualifier: involved_in
  review:
    summary: >-
      C. elegans sensory cilia are non-motile, and DYF-6 is required to build
      them: dyf-6 mutants have foreshortened amphid and phasmid ciliary endings.
      This is the most specific and organism-appropriate biological-process term
      for DYF-6.
    action: ACCEPT
    reason: >-
      Core biological process; the non-motile-cilium wording matches the worm
      sensory cilium and is directly supported by the mutant phenotype
      (PMID:16648645).
    supported_by:
    - reference_id: PMID:16648645
      supporting_text: the cilia of the amphid and phasmid dendritic endings are
        foreshortened
      reference_section_type: ABSTRACT
core_functions:
- description: >-
    DYF-6/IFT46 is a structural subunit of intraflagellar transport (IFT) complex
    B in ciliated sensory neurons. It localizes to the cilium and the ciliary base
    (basal body / transition-zone region), undergoes bidirectional IFT movement
    within the ciliated dendritic endings, and is required to build full-length
    non-motile sensory cilia β€” dyf-6 loss foreshortens the amphid and phasmid
    ciliary endings and produces the OSM-6 mislocalization pattern typical of IFT
    complex B mutants. It has no catalytic domain and acts as a structural/adaptor
    component of IFT-B; no informative molecular-function term is currently
    assignable (MF-dark; see knowledge_gaps).
  in_complex:
    id: GO:0030992
    label: intraciliary transport particle B
  directly_involved_in:
  - id: GO:0042073
    label: intraciliary transport
  - id: GO:1905515
    label: non-motile cilium assembly
  locations:
  - id: GO:0005929
    label: cilium
  - id: GO:0036064
    label: ciliary basal body
  supported_by:
  - reference_id: PMID:16648645
    supporting_text: Movement of DYF-6::GFP within the ciliated endings of the
      neurons indicates that DYF-6 is involved in IFT
    reference_section_type: ABSTRACT
  - reference_id: PMID:16648645
    supporting_text: the cilia of the amphid and phasmid dendritic endings are
      foreshortened
    reference_section_type: ABSTRACT
  - reference_id: PMID:28479320
    supporting_text: intraflagellar transport (IFT)-B complex abolishes dynein-2's
      ciliary localization
    reference_section_type: ABSTRACT
knowledge_gaps:
- gap_statement: >-
    The molecular function of DYF-6/IFT46 is undefined. It has a large disordered
    N-terminus and no recognizable catalytic domain, and no GO molecular-function
    term is assigned to it in C. elegans. Whether it acts as a cargo adaptor
    (Chlamydomonas IFT46 uses its N-terminal region to load outer dynein arms) or
    purely as an IFT-B scaffold, and which IFT-B subunit(s) it directly contacts
    in the worm, is not established.
  boundary: >-
    DYF-6 is firmly established as an IFT complex B subunit that localizes to the
    cilium and ciliary base, undergoes IFT movement, and is required to build
    full-length non-motile sensory cilia. In other organisms the conserved
    C-terminal IFT46_B_C domain mediates direct binding to IFT52 (and, via a
    ternary module, IFT88) to build the IFT-B1 core, while the N-terminal region
    binds the ODA16 cargo adaptor to load outer dynein arms β€” a role restricted
    to motile cilia and therefore not expected in the worm. What is missing is a
    GO molecular-function representation of this structural/scaffolding activity
    (the MF aspect cannot express "structural constituent of IFT particle B"), and
    the specific IFT-B contacts and any cargo-adaptor role of DYF-6 have not been
    demonstrated for the C. elegans protein.
  gap_kind:
  - ONTOLOGY
  - BIOLOGY
  dark_aspect: MF_DARK
  status: OPEN
  significance: >-
    IFT46 is a conserved core IFT-B protein; defining its molecular activity
    (scaffolding vs a specific cargo-adaptor interaction) is central to
    understanding IFT-B architecture and cargo selection in cilia.
  resolution: >-
    Map DYF-6's direct IFT-B interaction partners by biochemistry/structure;
    test for cargo-adaptor activity; introduce a molecular-function term for an
    IFT-particle structural/adaptor subunit activity.
  provenance:
  - reference_id: PMID:16648645
    supporting_text: DYF-6, the product of a complex locus, lacks known motifs,
      but orthologs are present in flies and mammals
    reference_section_type: ABSTRACT
  proposed_terms:
  - proposed_name: structural constituent of intraflagellar transport particle B
    proposed_definition: >-
      The action of a macromolecule that contributes to the structural integrity
      or scaffolding of an intraflagellar transport particle B (IFT-B) complex,
      as distinct from any catalytic or motor activity.
    justification: >-
      IFT-B subunits such as DYF-6/IFT46 have a well-defined cellular role (be
      part of, and maintain, the IFT-B particle) but no GO molecular-function term
      can currently express this structural/scaffolding activity, leaving them
      MF-dark despite complete BP/CC annotation.
    proposed_parent:
      id: GO:0005198
      label: structural molecule activity
- gap_statement: >-
    dyf-6 is a complex locus that, besides the short conserved IFT46 isoforms,
    produces a longer nematode-specific transcript fusing the dyf-6 ORF to the
    upstream gene F46F6.3; the biological role of this long product is
    uncharacterized. All functional and rescue data derive from the short
    isoforms, so whether the long isoform contributes anything beyond the
    conserved IFT-B function is unknown.
  boundary: >-
    The short (conserved) dyf-6 isoforms confer full dye-filling rescue and
    undergo IFT; the F46F6.3-fused long transcript is nematode-specific and has no
    assigned function, and existing alleles/reporters primarily report on the
    short forms.
  gap_kind:
  - BIOLOGY
  dark_aspect: RESIDUAL_SUBGAP
  status: OPEN
  significance: >-
    Establishing whether the nematode-specific long isoform has a distinct role
    would clarify whether the dyf-6 locus encodes more than the conserved IFT-B
    subunit function.
  resolution: >-
    Isoform-specific knockouts/rescue and expression analysis of the long
    dyf-6/F46F6.3 fusion product.
  provenance:
  - reference_id: PMID:16648645
    supporting_text: DYF-6, the product of a complex locus, lacks known motifs,
      but orthologs are present in flies and mammals
    reference_section_type: ABSTRACT
suggested_questions:
- question: >-
    Does C. elegans DYF-6/IFT46 act as a cargo adaptor within IFT-B (as
    Chlamydomonas IFT46 does for outer dynein arms), or purely as a structural
    scaffold, and which IFT-B subunits does it directly contact?
suggested_experiments:
- description: >-
    Affinity purification / cross-linking mass spectrometry and structural
    modeling of DYF-6 within the worm IFT-B complex to define its direct binding
    partners and position in the particle.
  experiment_type: interaction mapping
- description: >-
    Isoform-specific rescue and separation-of-function alleles (including
    N-terminal deletions) to test for a cargo-adaptor role and to determine
    whether the long dyf-6/F46F6.3 fusion isoform has any distinct function.
  experiment_type: genetics