WormMine: AnatomyTerm WBbt:0005755

• Definition: a rigid external coating which is secreted by the hypodermis, seam and some transitional epithelial cells which covers the outer body, the major openings into the body cavity from the exterior, and two large sensory bristles, the male spicules. The cuticle of the spicules, the hook, and of some portions of the spicule channels is especially rigid and is probably sclerotized. Similar sclerotic features are occasionally noted in the ventral surface of the male tail fan.
• Name: cuticle
• Primary Identifier: WBbt:0005755
• Synonym: exoskeleton

1 Children

Definition	Name	Synonym	Primary Identifier
Small linear ridges running along the lateral lines of the cuticle, lengthwise along the body; these thickenings lie in register with the seam cells which likely produce them and are present in the cuticle of the L1, dauer and adult stages, but not in other larval stages. The alae are suspected to provide better traction when the animal generates a bodywave. Since the animal usually lies on its side, the alae are in perfect position to catch against the substrate.	cuticular ala	alae	WBbt:0007832

0 Expression Clusters

24 Expression Patterns

• Primary Identifier: Expr16218
• Pattern: MLT-8::NeonGreen was co-localized with the lysosome marker CPL-1::mCherry in hyp7 and seam syncytia, and was secreted to the cuticle outside of both syncytia.

• Primary Identifier: Expr16428
• Pattern: Transgenic worms expressing mCherry under the daf-42 promoter showed expression in seam cells between 48 and 60 HAE, which corresponds to the time of appearance of the dead L2d phenotype. At 60 HAE, we observed expression in the seam cell, hypodermis, and at the surface of the worm. At 72 HAE, ecdysis had begun, and some worms had detached their head or tail from the old L2d cuticle, and the DAF-42::GFP signal did not remain in the L2d cuticle but was localized with the dauer worm inside the L2d cuticle. Together, our results show that daf-42 is expressed during the L2d-to-dauer transition in seam cells and is secreted toward the surface of the dauer worm, such as the hypodermis or cuticle.

• Primary Identifier: Expr16366
• Pattern: We found that IDPA-3, IDPB-3, IDPC-1, FIPR-4, and NSPB-12 are expressed exclusively in association with the pharynx and overlap in their localization with the pharynx cuticle. Briefly, tagged IDPA-3 was enriched in the grinder, overlapping the CFW-stained component and lining of the terminal bulb cuticle. In addition, we observed enrichment of tagged IDPA-3 in the presumptive ECM that lies between the terminal bulb and the intestinal valve. Tagged IDPB-3 was expressed weakly and localized exclusively to the pm6 cells and material surrounding the CFW-stained grinder. Tagged IDPC-1 had a similar pattern to that of tagged ABU-14; associating with both the anterior and posterior components of the pharyngeal cuticle. However, tagged ABU-14 appears to localize adjacent to CFW-stained components whereas tagged IDPC-1 overlaps CFW-stained components. Tagged NSPB-12 localized to the anterior pharynx cuticle components exclusively, including that of the buccal cavity, flaps, and anterior channels. Tagged FIPR-4 localized to both anterior and posterior pharynx cuticle components (but not the grinder teeth proper) and the presumptive pharynx-intestinal valve ECM.

• Primary Identifier: Expr13498
• Pattern: GFP::MAI-1 (xmSi32) and MAI-1::mCherry (xmSi31) showed the same tissue localization. GFP::MAI-1 is expressed in specific somatic tissues in the nuclei and in the cytoplasm through specific tissues, including the cuticle, hypodermis, rectum, vulva and neurons. Although the expression pattern did not suggest a mitochondria network and MAI-1 does not carry a mitochondrial transport sequence, we tested whether MAI-1 was expressed in the mitochondria by incubating GFP:: MAI-1 with Mitotracker Red CMXRos and found no co-localization between the mitochondrial probe and GFP::MAI-1. These results demonstrate that MAI-1 is indeed not expressed in mitochondria but in the cytoplasm, as previously suggested Ichikawa et al. Additionally, we observed that GFP::MAI-1 is expressed in the nuclei.

• Primary Identifier: Expr2400
• Remark: Reporter gene fusion type not specified.
• Pattern: The expression of Ce-cpz-1::gfp in transgenic C. elegans was restricted to the cuticular region along the length of the worm throughout development, and in the pharynx of only L3, L4 and adults.

• Primary Identifier: Expr3818
• Remark: Reporter gene fusion type not specified.
• Pattern: Expressed in adult. Expressed in hypodermal cells. No fluorescent signal could be discerned in embryos present in cln-3.2::GFP transgenic adults, in newly laid eggs from transgenic adults or in transgenic larvae. Adult cln-3.2::GFP transgenic hermaphrodite and male nematodes displayed fluorescence in the hypoderm visible as multiple thread-like patterns with regular interruptions and mild belt-like patterns running alongside of the body of the animal, both clearly distinguishable from autofluorescence. Punctate fluorescence in the pharynx and faint fluorescence in cells lining the cuticle of the head could be observed.

• Primary Identifier: Expr16367
• Pattern: We found that IDPA-3, IDPB-3, IDPC-1, FIPR-4, and NSPB-12 are expressed exclusively in association with the pharynx and overlap in their localization with the pharynx cuticle. Briefly, tagged IDPA-3 was enriched in the grinder, overlapping the CFW-stained component and lining of the terminal bulb cuticle. In addition, we observed enrichment of tagged IDPA-3 in the presumptive ECM that lies between the terminal bulb and the intestinal valve. Tagged IDPB-3 was expressed weakly and localized exclusively to the pm6 cells and material surrounding the CFW-stained grinder. Tagged IDPC-1 had a similar pattern to that of tagged ABU-14; associating with both the anterior and posterior components of the pharyngeal cuticle. However, tagged ABU-14 appears to localize adjacent to CFW-stained components whereas tagged IDPC-1 overlaps CFW-stained components. Tagged NSPB-12 localized to the anterior pharynx cuticle components exclusively, including that of the buccal cavity, flaps, and anterior channels. Tagged FIPR-4 localized to both anterior and posterior pharynx cuticle components (but not the grinder teeth proper) and the presumptive pharynx-intestinal valve ECM.

• Primary Identifier: Expr16368
• Pattern: We found that IDPA-3, IDPB-3, IDPC-1, FIPR-4, and NSPB-12 are expressed exclusively in association with the pharynx and overlap in their localization with the pharynx cuticle. Briefly, tagged IDPA-3 was enriched in the grinder, overlapping the CFW-stained component and lining of the terminal bulb cuticle. In addition, we observed enrichment of tagged IDPA-3 in the presumptive ECM that lies between the terminal bulb and the intestinal valve. Tagged IDPB-3 was expressed weakly and localized exclusively to the pm6 cells and material surrounding the CFW-stained grinder. Tagged IDPC-1 had a similar pattern to that of tagged ABU-14; associating with both the anterior and posterior components of the pharyngeal cuticle. However, tagged ABU-14 appears to localize adjacent to CFW-stained components whereas tagged IDPC-1 overlaps CFW-stained components. Tagged NSPB-12 localized to the anterior pharynx cuticle components exclusively, including that of the buccal cavity, flaps, and anterior channels. Tagged FIPR-4 localized to both anterior and posterior pharynx cuticle components (but not the grinder teeth proper) and the presumptive pharynx-intestinal valve ECM.

• Primary Identifier: Expr16369
• Pattern: We found that IDPA-3, IDPB-3, IDPC-1, FIPR-4, and NSPB-12 are expressed exclusively in association with the pharynx and overlap in their localization with the pharynx cuticle. Briefly, tagged IDPA-3 was enriched in the grinder, overlapping the CFW-stained component and lining of the terminal bulb cuticle. In addition, we observed enrichment of tagged IDPA-3 in the presumptive ECM that lies between the terminal bulb and the intestinal valve. Tagged IDPB-3 was expressed weakly and localized exclusively to the pm6 cells and material surrounding the CFW-stained grinder. Tagged IDPC-1 had a similar pattern to that of tagged ABU-14; associating with both the anterior and posterior components of the pharyngeal cuticle. However, tagged ABU-14 appears to localize adjacent to CFW-stained components whereas tagged IDPC-1 overlaps CFW-stained components. Tagged NSPB-12 localized to the anterior pharynx cuticle components exclusively, including that of the buccal cavity, flaps, and anterior channels. Tagged FIPR-4 localized to both anterior and posterior pharynx cuticle components (but not the grinder teeth proper) and the presumptive pharynx-intestinal valve ECM.

• Primary Identifier: Expr16370
• Pattern: We found that IDPA-3, IDPB-3, IDPC-1, FIPR-4, and NSPB-12 are expressed exclusively in association with the pharynx and overlap in their localization with the pharynx cuticle. Briefly, tagged IDPA-3 was enriched in the grinder, overlapping the CFW-stained component and lining of the terminal bulb cuticle. In addition, we observed enrichment of tagged IDPA-3 in the presumptive ECM that lies between the terminal bulb and the intestinal valve. Tagged IDPB-3 was expressed weakly and localized exclusively to the pm6 cells and material surrounding the CFW-stained grinder. Tagged IDPC-1 had a similar pattern to that of tagged ABU-14; associating with both the anterior and posterior components of the pharyngeal cuticle. However, tagged ABU-14 appears to localize adjacent to CFW-stained components whereas tagged IDPC-1 overlaps CFW-stained components. Tagged NSPB-12 localized to the anterior pharynx cuticle components exclusively, including that of the buccal cavity, flaps, and anterior channels. Tagged FIPR-4 localized to both anterior and posterior pharynx cuticle components (but not the grinder teeth proper) and the presumptive pharynx-intestinal valve ECM.

• Primary Identifier: Expr7814
• Remark: Picture: Fig. 3A.
• Pattern: Throughout development DPY-7 localises to the furrows that define the annuli in the wild type cuticle.

• Primary Identifier: Expr2933
• Pattern: SRP-2 expression is evident in numerous cells of the C. elegans embryo in the early stages of development (36-100-cell stage). However, in the later stages of embryonic development, expression is confined to a few cells of the anterior hypoderm and the eggshell. In the L1 and L2 larvae, expression is seen in a number of hypodermal (possibly hyp 1, 2, and 3) cells near the buccal cavity. Expression is also seen in the sensory support (sheath or socket) cells of several sensilla. Strong expression is visible in the posterior intestinal cells, seam cells, and the body hypoderm. In the adult hermaphrodite, strong punctate expression is seen in the hypodermal cells surrounding the anterior and posterior bulb of the pharynx. Expression was also seen in hyp 7 cells near the vulval opening. A strong striated staining pattern was seen in what appears to be the fibrous organelles that link muscle/hypoderm/cuticle. GFP expression was also observed in the phasmid (PHA and PHB) neurons, which was confirmed by co-staining with the amphid neuron-specific dye, DiI. The overall SRP-2::GFP expression pattern was similar in the hermaphrodites and males.

• Primary Identifier: Expr3899
• Pattern: Expressed in head/tail neurons, ventral nerve cord, alae, gut cells, vm1 and vulval epithelial cells

• Primary Identifier: Expr3126
• Pattern: The NAS-37::GFP fusion protein was found in the cuticle, with high concentration in the anterior of the worm overlying Hyp5. It was also concentrated in the alae, vulval cuticle and cuticle of the rectum. In addition, the protein was abundantly expressed in the excretory duct cell. The protein began to appear in the anterior cuticle 4 hours before each molt, consistent with the transcriptional GFP fusion results (see Expr3124 and Expr3125). Despite high levels of transcription of nas-37 in hypodermal cells, high levels of protein accumulation were not observed in these cells. GFP fluorescence was also observed in shed cuticles, demonstrating that the fusion protein is indeed secreted and deposited into the old cuticle.

• Primary Identifier: Expr11038
• Pattern: A transcriptional GFP reporter driven by the putative srap-1 promoter is detected in a variety of tissues including the major hypodermis and seam cells. Notable expression of this reporter is also observed in the tissues of the developing vulva and central nervous system. Expression of SRAP-1::GFP translational fusion is detected at the surface of the newly synthesized cuticle coincident with the molt. Significant accumulation of SRAP-1::GFP is also detected at the seam syncytium in the adult stage.

• Primary Identifier: Expr3897
• Pattern: Expressed in alae, head/tail neurons.

• Primary Identifier: Expr1776
• Remark: No GO_term assigned.
• Pattern: A distinct expression pattern in the hypodermal and cuticular regions of all stages was obtained from the Ce-cpl-1 translational fusion construct. Expression of the CPL-1::GFP fusion protein was also observed along the length of the pharyngeal lining in L1-L4 stages, but was restricted to the posterior bulb of the pharynx in adult worms. Furthermore, robust expression was observed in the eggshells surrounding the embryos inside the hermaphrodite and in the regions of the vulva, uterus, and spermatheca. In contrast to the transcriptional reporter construct there was no CPL-1::GFP expression in the gut.
• Subcellular Localization: GFP was detected in the eggshells of laid eggs in different stages of development.

• Primary Identifier: Expr2456
• Remark: The widespread adult cuticle expression pattern is remarkably similar to that previously described for rabbit polyclonal antibodies raised against total adult cuticles and probed against the surface of live worms, supporting the prediction that the GFP-tagged COL-19 expression may reflect the endogenous COL-19 expression pattern. Additionally, a recent study analyzing the localization of a Ty-tagged DPY-13 transgene displayed a remarkably similar annuli expression pattern to that described here for COL-19::GFP.
• Pattern: COL-19::GFP expression was also examined in dauer stage larvae. After allowing TP12:kaIs(col-19::gfp) strain to starve, a large population of dauer larvae were obtained, examined, and found not to express the COL-19::GFP protein in their cuticles. Likewise, no annuli or ala-specific GFP expression was detected in temperatureinduced dauers of a daf-2(e1370) mutant allele strain crossed with TP12:kaIs(col-19::gfp) males. COL-19::GFP is expressed exclusively in the adult cuticle, with a distinctive spatial expression pattern. The tagged protein is expressed in both forms of hypodermally derived cuticle, specifically, the matrix of the circumferential transverse annuli and the tri-laminate lateral alae. This dominant cuticle matrix expression was similar between adult hermaphrodites and males. In addition, the hermaphrodite-specific vulva fluoresced strongly, as did the male-specific tail structures, including the cuticular ray and fan structures.
• Subcellular Localization: COL-19::GFP was expressed in a transverse banding pattern of approximately 1.2 um in thickness, consistent with the annuli, and was absent from the annular furrows.

• Primary Identifier: Expr2362
• Pattern: GFP fluorescence was faint, but clearly observable on the surface of the larval and adult animal, presumably in the cuticle. This reporter gene did not rescue the Lon phenotype of lon-3 mutants, but instead caused an adult Rol (Roller) phenotype both in lon-3 mutants and in N2 wild type. These results suggest the cuticular localization of the reporter-gene product.

• Primary Identifier: Expr12489
• Pattern: abu-14.abu-14::sfGFP fluorescence localizes to the pharyngeal corpus and buccal cavity cuticle. Green fluorescence is seen in both the L4 and adult pharyngeal grinders, in the pharyngeal cuticle, and in the buccal cap.

• Primary Identifier: Expr16531
• Pattern: Endogenously tagged BLI collagens localize to puncta corresponding to cuticle struts. BLI-1::mNG fusion proteins were expressed exclusively in the late L4 epidermis and in the nascent and mature adult cuticle. Within the cuticle, BLI-1::mNG was localized to periodically repeating puncta of 0.25-0.3μm in diameter in cir- cumferential rows that extended from the dorsal or ventral midlines to the lateral alae. Based on their adult-specific expression and distribution, such puncta correspond to the cuticle struts originally observed in phase contrast and in TEM16, and are consistent with localization of an extrachromosomal transgenic BLI-1::GFP reporter31,33. BLI-1::mNG animals also expressed some diffuse mNG signal in the underlying epidermis beginning in the mid-L4 stage, presumably corresponding to BLI-1::mNG in the secretory pathway. XFP knock-ins within the BLI-1 N- and C-terminal domains (ju1789 and ju1791 respectively) displayed similar localization (Fig. S2a, b); most analysis below focused on the ju1789 N-terminal mNG knock-in in 1-2 day old adults.

• Primary Identifier: Expr16532
• Pattern: Endogenously tagged BLI collagens localize to puncta corresponding to cuticle struts. To determine the localization of BLI-2 we examined a BLI-2::mNG C-terminal knock-in (syb3293). BLI-2::mNG localized to adult strut puncta in a pattern essentially indistinguishable from that of BLI- 1::mNG. Additionally BLI-2::mNG localized to the lateral alae and seam-derived cuticle and to rings surrounding sensilla. BLI-2::mNG precisely colocalized with a BLI-1::wrmScarlet marker (juEx8105) in strut puncta, but not in the lateral alae.

• Primary Identifier: Expr16533
• Pattern: Endogenously tagged BLI collagens localize to puncta corresponding to cuticle struts. A BLI-6::mNG C-terminal knock-in (ju1914) localized to strut-like puncta. Unlike BLI-1::mNG and BLI-2::mNG, which tend to be more prominent in furrow-flanking strut rows, BLI-6::mNG was most prominent in the central row of struts in each annulus, confirmed by colocalization with BLI-1::wrmScarlet. Like BLI-2::mNG, BLI- 6::mNG localized to adult alae, seen as three very bright parallel ridges as well as more diffuse localization adjacent to the alae proper. BLI- 6::mNG additionally localized to cuticle furrows, and showed diffuse localization within the cuticle, as well as to larger foci in the deeper cuticle as well as to rings surrounding sensilla and the vulva.

• Primary Identifier: Expr16156
• Pattern: The same antibodies gave immunostaining patterns on C. elegans dauer larvae cuticles.

0 Life Stages

2 Parents

Definition	Name	Synonym	Primary Identifier
lies within the body wall, in close relation to nervous system and excretory system.	epithelial system		WBbt:0005730
Anatomical structures that exist outside of cells; non-cellular.	extracellular component		WBbt:0005732