WormMine: AnatomyTerm WBbt:0004939

• Definition: Male specific cephalic neurons (programmed cell death in hermaphrodite embryo) open to outside, possible function in male chemotaxis toward hermaphrodite.
• Name: CEMVR
• Primary Identifier: WBbt:0004939
• Synonym: lineage name: ABprpaapapp

1 Children

Definition	Name	Synonym	Primary Identifier
nucleus of pedigree ABprpaapapp	ABprpaapapp nucleus		WBbt:0001561

0 Expression Clusters

51 Expression Patterns

• Primary Identifier: Expr4254
• Remark: PKD-2::GFP transgenes fully rescue the male mating defect of pkd-2(sy606) null mutants.
• Reporter Gene: [pkd-2::gfp] translational fusion.
• Pattern: pkd-2 is expressed in a subset of male-specific sensory neurons: four CEM neurons in the head, eight pairs of Ray B neurons (1-5 and 7-9, excluding R6B), and the HOB hook neuron in the tail.
• Subcellular Localization: Each neuron that expresses PKD-2 has an exposed cilium at the distal ending of the dendrite. In these neurons, PKD-2::GFP localizes predominantly to cell bodies and cilia, and is also observed in small dendritic and axonal puncta. Within the ciliary region, PKD-2::GFP is distributed throughout the cilium, but primarily concentrated at the ciliary base, which corresponds to the distal-most dendrite and transition zone regions.

• Primary Identifier: Marker27
• Remark: Picture: Figure 3A.
• Pattern: Expressed in male specific sensory neurons.

• Primary Identifier: Expr12393
• Pattern: Pcil-7::GFP was expressed in CEM, RnB, HOB, and IL2 neurons.
• Subcellular Localization: The translational and functional CIL-7::GFP reporter localized to cell bodies (excluding nuclei), dendrites, axons, and the cilia and ciliary bases of the extracellular vesicle (EV)-releasing sensory neurons -CEM, RnB, HOB, and IL2.

• Primary Identifier: Expr9983
• Pattern: Pnphp-2::GFP was expressed in the ciliated sensory nervous system throughout development. In the adult, expression was evident in both hermaphrodite and male ciliated sensory neurons, including amphid, phasmid, and IL2 neurons. nphp-2 was also expressed in male specific ciliated sensory nervous system, including the CEM, RnB, and HOB neurons. Expression in the internal oxygen sensor neuron PQR was visible in the hermaphrodite, but could not be distinguished from the tail sensilla in the male.

• Primary Identifier: Expr3263
• Pattern: Pnphp-4::GFP is expressed in the amphid neurons in the head and the male-specific CEM neurons. Pnphp-4::GFP is expressed in both pairs of phasmids, the male-specific HOB and RnB neurons.

• Primary Identifier: Expr3720
• Pattern: In C. elegans males, nph-4::GFP expression in the head was seen in the amphid neurons, the outer labial neurons, and a cell group that could be compatible with the male-specific CEM neurons. In the male tail, nph-4::GFP expression was seen in the lumbar and cloacal ganglia. nph-4 expression in hermaphrodites was found in ciliated amphid neurons, the outer labial neurons, and the phasmid neurons in the tail. Developmental regulation of expression for nph-4::GFP was found to be similar to nph-1, appearing at the 1.5-fold stage and lasting through adulthood.

• Primary Identifier: Expr9653
• Pattern: CCPP-1::GFP was neuronally expressed in developing embryos (not shown) through adulthood in amphid and IL2 ciliated sensory neurons of the core nervous system, which males and hermaphrodites have in common. In the male-specific nervous system, CCPP-1::GFP was coexpressed with pkd-2 in CEM head neurons and the RnB and HOB neurons in the tail. CCPP-1::GFP was also expressed in some unidentified neurons and the gubernacular erector and retractor muscles in the male tail. CCPP-1::GFP was local- ized diffusely throughout neurons, including cilia, but excluded from the nucleus. CCPP-1::GFP was neuronally expressed in developing embryos (not shown) through adulthood in amphid and IL2 ciliated sensory neurons of the core nervous system, which males and hermaphrodites have in common. In the male-specific nervous system, CCPP-1::GFP was coexpressed with pkd-2 in CEM head neurons and the RnB and HOB neurons in the tail. CCPP-1::GFP was also expressed in some unidentified neurons and the gubernacular erector and retractor muscles in the male tail. CCPP-1::GFP was localized diffusely throughout neurons, including cilia, but excluded from the nucleus.

• Primary Identifier: Expr7848
• Remark: Picture: Fig. S1.
• Pattern: A ceh-30::gfp genomic construct rescued ceh-30(n4289{Delta}) for CEM survival and caused GFP expression in the nuclei of many neurons in the 3-fold embryo and in as many as a dozen neurons in larvae. GFP expression was not observed in the CEMs of male larvae, indicating that any ceh-30 expression in the CEM neurons is likely to be weak or transient. The ventral CEM neurons of 1.5-fold stage masculinized embryos, which can be identified by their positions within the embryo, expressed ceh-30::gfp. Expression in the dorsal CEMs cannot be as readily examined. Thus, ceh-30 is expressed transiently in embryonic CEMs.
• Subcellular Localization: GFP expression in the nuclei of many neurons in the 3-fold embryo.

• Primary Identifier: Expr15981
• Pattern: ENPP-1::mScarlet showed a speckled presence in the EVNs, with accumulations in the cilia of the CEM and shared IL2 neurons in the head and ray RnB neurons in the male tail. ENPP-1::mScarlet was released in environmental EVs from both CEMs and IL2s.

• Primary Identifier: Expr7849
• Remark: Picture: Fig. 4. Reporter gene fusion type not specified.
• Pattern: The CEH-30::GFP fusion is expressed widely during embryonic development, and the expression is especially robust in the anterior half of the embryos. CEH-30::GFP expression continues into the larval and adult stages, albeit at greatly reduced levels. In hermaphrodite smIs54 embryos, CEH-30::GFP was never seen in ventral CEMs (0/15 embryos). In contrast, in tra-1(n1099); smIs54 XX embryos, which are completely masculinized and virtually identical to male embryos, CEH-30::GFP was observed in all ventral CEMs (15/15 embryos), suggesting that CEH-30 is expressed in developing male CEM neurons.

• Primary Identifier: Expr7823
• Remark: Picture: Fig. 4.
• Subcellular Localization: OSM-9::GFP is localized to the cell body, but not cilia, of the polycystin neurons (CEM, HOB, RnB, where n = 1 to 9 except R6B)

• Primary Identifier: Expr3018
• Pattern: Expressed in AIN, AWA, BAG, HSN, URX. Faint or variable expression observed in an extra pair of cells in the tail. Male specific expression in Rays 5, 7, 9, CEM.

• Primary Identifier: Expr507
• Remark: Included upstream of gfp in the fusion gene was 5.1 kb of osm-6 genomic sequence, extending from 2.4 kb upstream of the 5' end of the osm-6 cDNA through the first 10 exons and all introns into the last exon, such that only the last 13 codons, including the translational stop, of OSM-6 were lacking. Transgenic Marker: rol-6(su1006) late embryo(author) = fully-elongated embryo(curator).
• Pattern: Expressed in 56 cells in the hermaphrodite: the 24 amphid neurons, twelve inner labial neurons, six outer labial neurons, four CEP cells, two ADE cells, four phasmid cells in all stages from late embryo through adult; AQR and PQR at L1 through adult PDE cells at L2 through adult. Additional expression in male-specific cells: CEM cells and male tail sensory cells.

• Primary Identifier: Expr9260
• Pattern: In the head of both males and hermaphrodites, the expression of the Ptbb-4::YFP transcriptional reporter was observed in many amphid neurons, likely ADL, AFD, ASE, ASG, ASH, ASJ, ASK, ASI, AWA, AWB, and AWC as judged by co-labeling with lipophilic dyes and/or cellular morphology. Other head neurons expressing Ptbb-4::YFP include the cephalic CEP neurons, the outer labial quadrant OLQ neurons, and the mechanosensory FLP neurons. In the midbody and tail, Ptbb-4::YFP expression was found in the PDE postdeirid neurons, PQR (a sensory neuron whose cilium is exposed to the pseudocoelomic fluid), and the phasmid neurons PHA and PHB. Ptbb-4::YFP was also expressed in all 18 ray neurons in the male, consistent with a previous report (Portman and Emmons 2004). Because of abundant ray neuron expression, Ptbb-4::YFP expression in male phasmid and hook neurons could not be reliably confirmed, although expression in these cells is not unlikely. We also observed Ptbb-4::YFP expression in the hermaphrodite HSN motor neurons, the vulval muscles, and several unidentified nonsensory neurons. Expression was also occasionally apparent in the amphid sheath (AMsh) and socket (AMso) glial cells.

• Primary Identifier: Expr12716

• Primary Identifier: Expr12717

• Primary Identifier: Expr843
• Remark: Reporter gene fusion type not specified. To analyze ida-1 gene expression in males, the pida-1::GFP transgenic strain BL5715 was crossed with the him-8 (high incidence of males) mutant strain BW1663. cgc6469 mentioned that this reporter gene is transcriptional fusion.
• Pattern: The ida-1 gene promoter-driven GFP expression pattern in males differs from hermaphrodites, especially in the tail, where many more fluorescent cells are observed. Male larvae appear very similar to hermaphrodite larvae, the only difference being that the PVP neurons in the preanal ganglion showed higher GFP expression levels. It was concluded that all neurons expressing GFP in the hermaphrodite pharyngeal nerve ring and in the tail also express GFP in males. In adult males, additional fluorescence is seen in neurons anterior to the nerve ring, in the ventral cord and many more in the tail. The four additional GFP-expressing cells in the procorpus just anterior to the pharyngeal bulb have not been identified. Expression in the male-specific CEM neurons at the anterior end of the nerve ring appeared variable. The hermaphrodite-specific cells VC, HSN, and uv1 observed to express GFP are not found in males. Nonetheless, several cell bodies in the male ventral cord expressed GFP. Dorsally directed processes emanate from them, which identifies them as CAn, a set of eight motoneurons. As in hermaphrodites, PDE weakly expressed GFP. In the male tail, about 20 neurons express GFP. They include PHA, PHB, PHC, and PVP in the preanal ganglion, which are already seen in larvae. Some of the rays contain GFP-filled axons that extend all the way to the tip of the ray. Rays 2, 8, and 9 show the highest expression levels; axons of rays 1, 4, and 6 fluoresce more weakly. No GFP expression is observed in the spicules.

• Primary Identifier: Expr1315
• Pattern: ADL, ASH, ASK, PHA, PHB, distal tip cell, anchor cell, many male-specific neurons.

• Primary Identifier: Expr10037
• Pattern: daf-37 is expressed in the ASI and ASK sensory neurons, in the IL-2 interneurons, and in the male-specific CEM neurons. DAF-37 is strongly expressed in L1 and L2 stages but is less abundant in the L3, L4, and adult stages. We also detected DAF-37 in dauer larvae, although the level of expression was lower than in any other stages.
• Subcellular Localization: DAF-37 is localized to the cilia.

• Primary Identifier: Expr12727

• Primary Identifier: Expr8036
• Remark: Picture: Figure 2B. stam-1, also called pqn-19.
• Pattern: stam-1 is expressed in many tissues, including the pharyngeal intestinal valve, several head neurons, and phasmids in both males and hermaphrodites throughout development. In males, stam-1 expression is also observed in the gonad and sensory neurons in the tail. stam-1 and pkd-2 are clearly coexpressed in male-specific ciliated CEM, ray B-type (RnB), and hook HOB sensory neurons.

• Primary Identifier: Expr9327
• Subcellular Localization: PKD-2::DsRed2 localizes to cilia and cell bodies of ray, HOB, and CEM neurons. ATP-2::GFP clearly colocalizes with PKD-2::DsRed2 in CEM cilia. In rays and HOB, ciliary colocalization of ATP-2 and PKD-2 was ambiguous due to the expression of ATP-2::GFP in ray neurons, support cells, and hypodermis.

• Primary Identifier: Expr8949
• Remark: Picture: Fig. 1B to 1D.
• Pattern: Expression of both of these transgenes was observed specifically in the RnB, HOB and CEM neurons of the male, and nowhere in the hermaphrodite.
• Subcellular Localization: The CWP-5::YFP fusion protein was clearly enriched in the sensory cilia of these neurons and was also detectable in the soma.

• Primary Identifier: Expr2408
• Pattern: PKD-2::GFP4 is expressed in the male-specific sensory neurons of the head (the CEMs), rays, and hook (HOB).
• Subcellular Localization: KD-2(+)::GFP4 concentrates in sensory cilia and in a punctate pattern in neuronal cell bodies. Punctate pkd-2::gfp4 expression in ray neuronal cell bodies may be in RER or in Golgi. PKD-2::GFP4 accumulates in a ring around the ciliary transition zones, consistent with plasma membrane localization.

• Primary Identifier: Expr2409
• Remark: The overall pattern of alpha-PKD-2 antibody staining is consistent with the expression of the pkd-2::gfp4 fusion gene, although there are a few differences. First, alpha -PKD-2 antibodies label only male-specific neurons, whereas pkd-2::gfp4 is additionally expressed in non-sex-, non-stage-specific nerve ring neurons and occasionally in ray structural cells. Second, alpha-PKD-2 antibodies also recognize axons of male-specific sensory neurons, whereas pkd-2::gfp4 is only weakly expressed in axons. pkd-2(sy606 deletion) mutant animals exhibited no immunoreactivity, demonstrating the specificity of our alpha-PKD-2 polyclonal antibody.
• Pattern: The alpha-PKD-2 antibodies stained only male-specific sensory neurons in wild-type animals. Based on cell body positions, the PKD-2 protein is detected in the head CEMs, ray (except Ray 6), and hook HOB neurons of wild-type males.
• Subcellular Localization: The alpha-PKD-2 antibodies labeled the entire neuron (cilia, dendrite, cell body, and axon). Cytoplasmic, nonnuclear staining in cell bodies and dendrites is punctate, suggesting that one pool of PKD-2 is located in intracellular membranes of the ER, Golgi apparatus, and/or transport vesicles while another is found in sensory cilia.

• Primary Identifier: Expr3208
• Pattern: lov-1 and atp-2 are coexpressed in the tail ray B neurons and HOB hook neuron as well as the male-specific CEM head neurons.
• Subcellular Localization: ATP-2::GFP is found in a broad range of tissue types, making it difficult to ascertain ciliary localization. To look more carefully at ATP-2 subcellular localization, we examined colocalization of ATP-2 and PKD-2. LOV-1 and PKD-2 exhibit identical subcellular localization patterns. PKD-2::DsRed2 localizes to cilia and cell bodies of ray, HOB, and CEM neurons. ATP-2::GFP clearly colocalizes with PKD-2::DsRed2 in CEM cilia. In rays and HOB, ciliary colocalization of ATP-2 and PKD-2 was ambiguous due to the expression of ATP-2::GFP in ray neurons, support cells, and hypodermis.

• Primary Identifier: Expr3490
• Subcellular Localization: In the wild-type, PKD-2::GFP localizes throughout neuronal cell bodies and in small, discrete puncta in cilia of male-specific CEM, RnB, and HOB neurons. In wild-type rays, PKD-2::GFP normally localizes to ciliated endings on dendrites of RnB neurons. In the head of a wild-type male, PKD-2::GFP normally localizes to the ciliary transition zone and small, finger-like ciliary projections emanating from the transition zone of the four CEM neurons.

• Primary Identifier: Expr9657
• Pattern: klp-6 is expressed in the male-specific sensory neurons of the rays (RnBs, n = 1-9 but not 6), hook (HOB), and head (CEMs). klp-6 is also expressed in the IL2 head sensory neurons of males and hermaphrodites.
• Subcellular Localization: Genomic and cDNA fusions (gKLP-6::GFP and cKLP-6::GFP, respectively) are expressed in an identical cell-specific pattern and give equivalent localization patterns. KLP-6::GFP is observed throughout the neuron in the cell body, axon, dendrite, and cilium, and it is excluded from the nucleus.

• Primary Identifier: Expr3262
• Pattern: Pnphp-1::GFP is expressed in both male and hermaphrodite. Pnphp-1::GFP is expressed in the amphid neurons in the head and the male-specific CEM neurons. Pnphp-1::GFP is also expressed in the male and hermaphrodite tail. Pnphp-1::GFP is expressed in the ADL amphid neurons, both pairs of phasmids, the male-specific HOB and RnB neurons, and a subset of the postcloacal sensillae neurons.

• Primary Identifier: Expr1148
• Remark: Reporter gene fusion type not specified.
• Pattern: A 6.7-kb fusion (LOV-1::GFP1) and a 15.4-kb fusion lacking only 58 amino acids (LOV-1::GFP5) direct expression in male-specific sensory neurons. Shorter versions of lov-1::gfp1 are not expressed in the same set of male-specific neurons, nor do they act as dominant negatives. lov-1::gfp1 and lov-1::gfp5 are expressed in male-sensory neurons, specifically the CEM neurons, the hook neuron HOB, and the sensory ray neurons. Expression begins during late L4 lethargus and peaks in the adult male.
• Subcellular Localization: LOV-1::GFP1 is localized at high levels in the cell body (punctate cytoplasmic expression) and ciliated endings, whereas LOV-1::GFP5 is found exclusively in cell bodies.

0 Life Stages

4 Parents

Definition	Name	Synonym	Primary Identifier
Neuron class of four male-specific neurons associated with cephalic sensilla.	CEM		WBbt:0005246
cell that exists in and is part of an embryo.	embryonic cell		WBbt:0007028
embryonic cell	ABprpaapap		WBbt:0006105
embryonic cell	ABprpaapapp		WBbt:0006292