C. elegans detects volatile odorants through amphid sensory neurons that express diverse G protein-coupled receptors (GPCRs), many of which remain functionally uncharacterized. We evaluated the candidate chemosensory GPCR srd-44 using the srd-44(ok1831) mutant strain in chemotaxis assays with the defined odorant diacetyl and the complex natural attractant Carolina mantle (Philomycus carolinianus) slug mucus. Wild-type and mutant animals exhibited similar chemotaxis responses to both stimuli. Under these conditions, these results do not support a role for srd-44 in chemotaxis toward diacetyl or Carolina mantle slug mucus. These findings contribute to the functional evaluation of an orphan chemosensory GPCR in C. elegans.
","acknowledgements":"We acknowledge wormbase (now Alliance of Genome Resources), the Caenorhabditis Genetics Center, and the C. elegans Gene Knockout Project at the Oklahoma Medical Research Foundation (part of the International C. elegans Gene Knockout Consortium) for their contributions that make research such as ours possible.
","authors":[{"affiliations":["Goucher College, Towson, MD, US"],"departments":["Biological Sciences"],"credit":["formalAnalysis","fundingAcquisition","resources","supervision","writing_originalDraft","visualization","writing_reviewEditing"],"email":"jay.garaycochea@goucher.edu","firstName":"Jay","lastName":"Garaycochea","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0009-0007-5724-2667"},{"affiliations":["Goucher College, Towson, MD, US"],"departments":["Biological Sciences"],"credit":["conceptualization","investigation","methodology"],"email":"milo.roth@Goucher.edu","firstName":"Milo","lastName":"Roth","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"Funding for the project was provided by the The Clara W. Claasen '25 Scholarship and start-up funds from Goucher College.
","image":{"url":"https://portal.micropublication.org/uploads/fb53c91b4476a7c44f2c2c9994927c34.png"},"imageCaption":"A. Chemotaxis index (CI) of wild-type (N2) and srd-44(ok1831) (RB1526) animals toward diacetyl (1:100 v/v in ethanol), Carolina mantle slug mucus extract, Isoamyl Alcohol (1:100 v/v in ethanol), 1-Octanol (1:10 v/v in ethanol), and ethanol control after 60 minutes. CI was calculated as (T − C) / (T + C), where T represents animals in the test quadrants and C represents animals in ethanol control quadrants. Data are shown as individual biological replicates with mean ± SD. Statistical analysis was performed using two-way ANOVA (genotype x stimulus) followed by Tukey’s multiple comparisons test. B. Chemotaxis index (CI) of wild-type (N2) and srd-44(ok1831) (RB1526) animals towards diacetyl concentrations ranging from 1.14 × 10⁻⁵ M to 1.14 M after 60 minutes. Data shown as mean ± SD from 3-10 independent assay plates per concentration. No consistent differences in chemotaxis responses were observed between wild-type and RB1526 animals across the concentration range tested.
","imageTitle":"Loss of srd-44 is not associated with reduced chemotaxis to various odorants
","methods":"N2 and RB1526 (srd-44(ok1831)) strains were obtained from the Caenorhabditis Genetics Center (CGC). The srd-44(ok1831) allele contains a 2064 bp deletion withing the srd-44 (F17A2.8) locus on chromosome X and is predicted to cause transcript ablation of the primary srd-44 transcript. This allele was generated by the C. elegans Gene Knockout Project at the Oklahoma Medical Research Foundation, which was part of the International C. elegans Gene Knockout Consortium. The srd-44(ok1831) allele deletion was confirmed through a primer pair of inner left (cttgatcagtcgctctcgtg) and inner right (cgcaaccattttggagagac) following six backcrossings. C. elegans strains were maintained using standard nematode culture protocols as previously described (Stiernagle, 2006). Briefly, an OP50 E. coli lawn was grown on NGM-coated 60 mm petri plates. Picked eggs were transferred from freshly starved stock plates, incubated at 20°C for 2-3 days, then noted for presence of synchronized L4 and young adults. Worms were collected and pelleted using an M9 wash buffer before being transferred to a prepared testing plate.
Chemotaxis assay was adopted from Margie et al. (2013) by utilizing four quadrant divisions with two test and two control quadrants in a 10cm petri plate. Pelleted worms were placed in the center and allowed to absorb into the agar surface for 5 minutes. Testing compounds, mixed with 0.5M sodium azide, were applied to designated areas. After one hour at room temperature, plates were placed into a 4°C cooler, then scored for worms that travelled 0.5 cm from the point of origin. Chemotaxis Index was calculated using the following equation: Chemotaxis Index = (# Worms in Both Test Quadrants - Worms in Both Control Quadrants) / (Total # of Scored Worms).
Philomycus carolinianus (Carolina Mantle slug) was collected from the deciduous forests surrounding Goucher College in Towson, Maryland USA. Slugs were placed in a terrarium for 24 hours before being transferred to a terrarium containing autoclaved dirt, rocks, and woody material. Slugs were fed a diet of autoclaved carrots and lettuce, misted with autoclaved tap water via a Honeywell humidifier, and allowed to acclimate for 3 days before mucus was collected. Cotton tipped applicators were used to collect mucus, then swirled in an Eppendorf tube with 45 µl of ethanol and stored in a 4°C cooler before being transferred to a -80°C freezer. All slugs were returned to their native habitat at the end of the study.
Statistics were performed using Graphpad Prism software version 10.5.0 for windows. A two-way ANOVA was used to evaluate the effects of genotype and stimulus condition, followed by Tukey’s multiple comparisons to assess genotype differences within each condition. Descriptive statistics were used to calculate mean and standard deviation. The figure was created using grouped summary data.
Tukey's multiple comparisons test (Wildtype vs. RB1526) | Predicted (LS) Mean diff. | Adjusted P Value |
Diacetyl | -0.2448 to 0.2867 | 0.8733 |
Carolina Mantleslug | -0.4895 to 0.1697 | 0.3300 |
Isoamyl Alcohol | -0.4793 to 0.06227 | 0.1265 |
1-Octanol | -0.07980 to 0.5368 | 0.1408 |
Ethanol | -0.3069 to 0.3522 | 0.8895 |
OpenAI’s ChatGPT, version GPT-5.2, was used for editing language, clarity, and structure of the author’s original draft manuscript of the description and abstract. Title, figures, figure legend, and methods were author originated. Prompts requesting deep research into previous experimental findings, suggestions for references, and critique of conclusions were manually reviewed and verified by the authors. The authors accept full responsibility for the accuracy of all content in the final manuscript.
","reagents":"Strain | Genotype | Available From |
N2 | Caenorhabditis elegans | CGC |
srd-44(ok1831) | F17A2.8. Homozygous | CGC |
Species | Common Name | Location Acquired |
Philomycus carolinianus | Carolina Mantle slug | Woods surrounding Goucher College |
Reagent | Source | Description |
1-Octanol | Sigma Aldrich | Cat# 472328-100ML |
Diacetyl | TCI Chemicals | Cat# B0682-25ML |
Ethanol | Pharmco | Cat# 111000190-1Gal |
Isoamyl Alcohol | Sigma Aldrich | Cat# W205702-1KG-K |
C. elegans detects volatile odorants through amphid sensory neurons that express diverse G protein-coupled receptors (GPCRs), many of which remain functionally uncharacterized (Bargmann et al., 1993; Robertson & Thomas, 2006). Members of this large chemosensory GPCR family mediate responses to both defined odorants and complex biological compounds (Bargmann et al., 1993; Kim et al., 2009; Troemel et al., 1995). Identifying the ligands of orphaned GPCRs is a challenging and time-consuming task that requires a comprehensive approach integrating a multitude of techniques to understand the full range of GPCR responses (Jobe & Vijayan, 2024). One potential approach is using biological complex compounds with gas chromatography/mass spectrometry to screen for candidate metabolites (Gaerlan et al., 2025).
C. elegans coexist in natural environments with terrestrial invertebrates including isopods and gastropods such as slugs (Frézal & Félix, 2015). Slug mucus contains a mixture of proteins, glycoproteins, and small metabolites that act as volatile compounds and may serve as multimodal chemical cues (Ballard et al., 2021; Smith & Morin, 2002). Members of the large chemosensory GPCR family expressed in amphid neurons mediate responses to both defined odorants and complex biological compounds (Bargmann et al., 1993; Kim et al., 2009; Troemel et al., 1995). The properties of slug mucus position it as a candidate for screening odorant GPCRs in C. elegans.
srd-44 encodes a predicted seven-transmembrane receptor belonging to the serpentine receptor class delta (srd) family, a group of genes enriched in chemosensory neurons (Robertson & Thomas, 2006; Troemel et al., 1995; Vidal et al., 2018). A previous reporter-based study suggested expression of srd-44 in ASH amphid sensory neurons, although endogenous expression patterns remain incompletely defined (Wood & Ferkey, 2019). Despite its predicted classification as a chemosensory receptor, no ligand or defined behavioral function has been reported for srd-44. srd-44 was selected as a candidate for behavioral screening due to the putative expression in ASH sensory neuron allowing for interaction with volatile odorants.
Wild-type (N2) and RB1526 (srd-44(ok1831)) C. elegans were compared for chemotaxis toward diacetyl and Carolina mantle slug mucus (Figure panel A). No differences in chemotaxis were observed between genotypes for either stimulus under the conditions tested. To further evaluate responses to diacetyl, both strains were subsequently examined using diacetyl concentrations ranging from 1.14 × 10⁻⁵ M to 1.14 M (Figure panel B). Both genotypes exhibited similar concentration-dependent chemotaxis responses across the tested range, with no difference observed between wild-type and RB1526 animals.
","references":[{"reference":"Ballard KR, Klein AH, Hayes RA, Wang T, Cummins SF. 2021. The protein and volatile components of trail mucus in the Common Garden Snail, Cornu aspersum. PLoS One 16(5): e0251565.
","pubmedId":"34043643","doi":""},{"reference":"Bargmann CI, Hartwieg E, Horvitz HR. 1993. Odorant-selective genes and neurons mediate olfaction in C. elegans. Cell 74(3): 515-27.
","pubmedId":"8348618","doi":""},{"reference":"Frézal L, Félix MA. 2015. C. elegans outside the Petri dish. Elife 4: 10.7554/eLife.05849.
","pubmedId":"25822066","doi":""},{"reference":"Gaerlan M, Carrillo M, Ceva S, Chundi S, Diallo B, Fong JN, et al., O'Connell LA. 2025. Velvety tree ant extract is a chemotaxis repellent for C. elegans. MicroPubl Biol 2025: 10.17912/micropub.biology.001531.
","pubmedId":"40535527","doi":""},{"reference":"Jobe A, Vijayan R. 2024. Orphan G protein-coupled receptors: the ongoing search for a home. Front Pharmacol 15: 1349097.
","pubmedId":"38495099","doi":""},{"reference":"Kim K, Sato K, Shibuya M, Zeiger DM, Butcher RA, Ragains JR, et al., Sengupta P. 2009. Two chemoreceptors mediate developmental effects of dauer pheromone in C. elegans. Science 326(5955): 994-8.
","pubmedId":"19797623","doi":""},{"reference":"Margie O, Palmer C, Chin-Sang I. 2013. C. elegans chemotaxis assay. J Vis Exp(74): e50069.
","pubmedId":"23644543","doi":""},{"reference":"Robertson HM, Thomas JH. 2006. The putative chemoreceptor families of C. elegans. WormBook: 1-12.
","pubmedId":"18050473","doi":""},{"reference":"Sengupta P, Chou JH, Bargmann CI. 1996. odr-10 encodes a seven transmembrane domain olfactory receptor required for responses to the odorant diacetyl. Cell 84(6): 899-909.
","pubmedId":"8601313","doi":""},{"reference":"Smith AM, Morin MC. 2002. Biochemical differences between trail mucus and adhesive mucus from marsh periwinkle snails. Biol Bull 203(3): 338-46.
","pubmedId":"12480724","doi":""},{"reference":"Stiernagle T. 2006. Maintenance of C. elegans. WormBook: 1-11.
","pubmedId":"18050451","doi":""},{"reference":"Troemel ER, Chou JH, Dwyer ND, Colbert HA, Bargmann CI. 1995. Divergent seven transmembrane receptors are candidate chemosensory receptors in C. elegans. Cell 83(2): 207-18.
","pubmedId":"7585938","doi":""},{"reference":"Vidal B, Aghayeva U, Sun H, Wang C, Glenwinkel L, Bayer EA, Hobert O. 2018. An atlas of Caenorhabditis elegans chemoreceptor expression. PLoS Biol 16(1): e2004218.
","pubmedId":"29293491","doi":""},{"reference":"Wood J, Ferkey D. 2019. unc-42 regulates the expression of ASH terminal fate markers. MicroPubl Biol 2019: 10.17912/micropub.biology.000114.
","pubmedId":"32550468","doi":""}],"title":"Behavioral Screening of the Candidate Chemoreceptor Gene srd-44 Using Diacetyl and Carolina Mantle Slug Mucus in Caenorhabditis elegans
","reviews":[{"reviewer":{"displayName":"Erika Sorensen"},"openAcknowledgement":false,"status":{"submitted":true}}],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":null}]},{"id":"c84cd503-0a4e-43e7-a8a4-9fe0964359ad","decision":"revise","abstract":"C. elegans detect volatile odorants through amphid sensory neurons that express diverse G protein-coupled receptors (GPCRs), many of which remain functionally uncharacterized. We evaluated the candidate chemosensory GPCR srd-44 using the RB1526 mutant strain in chemotaxis assays against diacetyl and Carolina mantle (Philomycus carolinianus) slug mucus. Wild-type and mutant animals exhibited similar chemotaxis responses to both stimuli across a range of concentrations. Under these conditions, these results do not support a role for srd-44 in chemotaxis toward diacetyl or Carolina mantle slug mucus. These findings contribute to the functional evaluation of an orphan chemosensory GPCR in C. elegans.
","acknowledgements":"We acknowledge wormbase (now Alliance of Genome Resources), the Caenorhabditis Genetics Center, and the C. elegans Gene Knockout Project at the Oklahoma Medical Research Foundation (part of the International C. elegans Gene Knockout Consortium) for their contributions that make research such as ours possible.
","authors":[{"affiliations":["Goucher College, Towson, MD, US"],"departments":["Biological Sciences"],"credit":["formalAnalysis","fundingAcquisition","resources","supervision","writing_originalDraft","visualization","writing_reviewEditing"],"email":"jay.garaycochea@goucher.edu","firstName":"Jay","lastName":"Garaycochea","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0009-0007-5724-2667"},{"affiliations":["Goucher College, Towson, MD, US"],"departments":["Biological Sciences"],"credit":["conceptualization","investigation","methodology"],"email":"milo.roth@Goucher.edu","firstName":"Milo","lastName":"Roth","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"Funding for the project was provided by the The Clara W. Claasen '25 Scholarship and start-up funds from Goucher College.
","image":{"url":"https://portal.micropublication.org/uploads/bb779c490f393b058b95d24f29e21c30.png"},"imageCaption":"A. Chemotaxis index (CI) of wild-type (N2) and RB1526 (srd-44(ok1831)) animals toward diacetyl (1:100 v/v in ethanol), Carolina mantle slug mucus extract, Isoamyl Alcohol (1:100 v/v in ethanol), 1-Octanol (1:10 v/v in ethanol), and ethanol control after 60 minutes. CI was calculated as (T − C) / (T + C), where T represents animals in the test quadrants and C represents animals in ethanol control quadrants. Data are shown as individual biological replicates with mean ± SD. Statistical analysis was performed using two-way ANOVA (genotype x stimulus) followed by Tukey’s multiple comparisons test. B. CI of wild-type and RB1526 animals towards diacetyl concentrations ranging from 1.14 × 10⁻⁵ M to 1.14 M after 60 minutes. Data shown as mean ± SD from 3-10 biological replicates per concentration. C. CI of wild-type and RB1526 animals towards Carolina Mantle Slug mucus concentrations ranging from a 100 to 10-5 dilution from original sample after 60 minutes. Data shown as mean ± SD from 3-4 biological replicates per concentration. No consistent differences in chemotaxis responses were observed between wild-type and RB1526 animals across the concentration range tested.
","imageTitle":"Loss of srd-44 is not associated with reduced chemotaxis to various odorants
","methods":"N2 and RB1526 (srd-44(ok1831)) strains were provided by the CGC, which is funded by NIH Office of Research Infrastructure Programs (P40 OD010440). The srd-44(ok1831) allele contains a 2064 bp deletion withing the srd-44 (F17A2.8) locus on chromosome X and is predicted to cause transcript ablation of the primary srd-44 transcript (Alliance of Genome Resources Consortium, 2024; Sternberg et al., 2024). This allele was generated by the C. elegans Gene Knockout Project at the Oklahoma Medical Research Foundation, which was part of the International C. elegans Gene Knockout Consortium. The srd-44(ok1831) allele deletion was confirmed through a primer pair of inner left (cttgatcagtcgctctcgtg) and inner right (cgcaaccattttggagagac) following six backcrossings. C. elegans strains were maintained using standard nematode culture protocols as previously described (Stiernagle, 2006). Briefly, an OP50 E. coli lawn was grown on NGM-coated 60 mm petri plates. Picked eggs were transferred from freshly starved stock plates, incubated at 20°C for 2-3 days, then noted for presence of synchronized L4 and young adults. Worms were collected and pelleted using an M9 wash buffer before being transferred to a prepared testing plate.
Chemotaxis assay was adopted from Margie et al. (2013) by utilizing four quadrant divisions with two test and two control quadrants in a 10cm petri plate. 40-150 Pelleted worms were placed in the center and allowed to absorb into the agar surface for 5 minutes. Testing compounds, mixed with 0.5M sodium azide, were applied to designated areas. After one hour at room temperature, plates were placed into a 4°C cooler, then scored for worms that travelled 0.5 cm from the point of origin. Chemotaxis Index was calculated using the following equation: Chemotaxis Index = (# Worms in Both Test Quadrants - Worms in Both Control Quadrants) / (Total # of Scored Worms).
Philomycus carolinianus (Carolina Mantle slug) was collected from the deciduous forests surrounding Goucher College in Towson, Maryland USA. Slugs were placed in a terrarium for 24 hours before being transferred to a terrarium containing autoclaved dirt, rocks, and woody material. Slugs were fed a diet of autoclaved carrots and lettuce, misted with autoclaved tap water via a Honeywell humidifier, and allowed to acclimate for 3 days before mucus was collected. Cotton tipped applicators were used to collect mucus, then swirled in an Eppendorf tube with 45 µl of ethanol and stored in a 4°C cooler before being transferred to a -80°C freezer. Ethanol was selected as a solvent for the mucus to account for the effects of the solvent in behavioral assays. All slugs were returned to their native habitat at the end of the study.
Statistics were performed using Graphpad Prism software version 10.5.0 for windows. A two-way ANOVA was used to evaluate the effects of genotype and stimulus condition, followed by Tukey’s multiple comparisons to assess genotype differences within each condition. Descriptive statistics were used to calculate mean and standard deviation. The figure was created using grouped summary data.
OpenAI’s ChatGPT, version GPT-5.2, was used for editing language, clarity, and structure of the author’s original draft manuscript of the description and abstract. Title, figures, figure legend, and methods were author originated. Prompts requesting deep research into previous experimental findings, suggestions for references, and critique of conclusions were manually reviewed and verified by the authors. The authors accept full responsibility for the accuracy of all content in the final manuscript.
","reagents":"Strain | Genotype | Available From |
N2 | Caenorhabditis elegans | CGC |
RB1526 | srd-44(ok1831) | CGC |
Species | Common Name | Location Acquired |
Philomycus carolinianus | Carolina Mantle slug | Woods surrounding Goucher College |
Reagent | Source | Description |
1-Octanol, ≥99% (GC) | Sigma Aldrich | Cat# 472328-100ML |
Diacetyl, 98.0+% | TCI Chemicals | Cat# B0682-25ML |
Ethanol, 200 proof | Pharmco | Cat# 111000190-1Gal |
Isoamyl Alcohol, ≥98% (GC) | Sigma Aldrich | Cat# W205702-1KG-K |
Statistical Analysis
Tukey's multiple comparisons test (Wildtype vs. RB1526) | Predicted (LS) Mean diff. | Adjusted P Value |
Diacetyl | -0.2448 to 0.2867 | 0.8733 |
Carolina Mantleslug | -0.4895 to 0.1697 | 0.3300 |
Isoamyl Alcohol | -0.4793 to 0.06227 | 0.1265 |
1-Octanol | -0.07980 to 0.5368 | 0.1408 |
Ethanol | -0.3069 to 0.3522 | 0.8895 |
C. elegans detects volatile odorants through amphid sensory neurons that express diverse G protein-coupled receptors (GPCRs), many of which remain functionally uncharacterized(Bargmann et al., 1993; Robertson & Thomas, 2006). Members of this large chemosensory GPCR family mediate responses to both defined odorants and complex biological compounds(Bargmann et al., 1993; Kim et al., 2009; Troemel et al., 1995). Identifying the ligands of orphaned GPCRs is a challenging and time-consuming task that requires a comprehensive approach integrating a multitude of techniques to understand the full range of GPCR responses (Jobe & Vijayan, 2024). One potential approach is using biological complex compounds with gas chromatography/mass spectrometry to screen for candidate metabolites (Gaerlan et al., 2025).
C. elegans coexist in natural environments with terrestrial invertebrates including isopods and gastropods such as slugs (Frézal & Félix, 2015). Slug mucus contains a mixture of proteins, glycoproteins, and small metabolites that act as volatile compounds and may serve as multimodal chemical cues (Ballard et al., 2021; Smith & Morin, 2002). Members of the large chemosensory GPCR family expressed in amphid neurons mediate responses to both defined odorants and complex biological compounds (Bargmann et al., 1993; Kim et al., 2009; Troemel et al., 1995). The properties of slug mucus position it as a candidate for screening odorant GPCRs in C. elegans.
srd-44 encodes a predicted seven-transmembrane receptor belonging to the serpentine receptor class delta (srd) family, a group of genes enriched in chemosensory neurons(Robertson & Thomas, 2006; Troemel et al., 1995a; Vidal et al., 2018). A previous reporter-based study suggested expression of srd-44 in ASH amphid sensory neurons, although endogenous expression patterns remain incompletely defined (Wood & Ferkey, 2019). ASH neurons are implicated in avoidance response to nose touch and avoidance of noxious chemicals (Kaplan & Horvitz, 1993). Despite its predicted classification as a chemosensory receptor, no ligand or defined behavioral function has been reported for srd-44. srd-44 was selected as a candidate for behavioral screening due to the putative expression in ASH sensory neuron allowing for interaction with volatile odorants.
Wild-type (N2) and RB1526 (srd-44(ok1831))C. elegans were compared for chemotaxis towards the volatile attractant diacetyl and isoamyl alcohol, the volatile repellent 1-octanol, and the undefined cue Carolina mantle slug mucus, with ethanol as the negative control (Figure 1A). No differences in chemotaxis were observed between genotypes for either stimulus under the conditions tested. To further evaluate responses to diacetyl, both strains were subsequently examined using diacetyl concentrations ranging from 1.14 × 10⁻⁵ M to 1.14 M (Figure 1B) and Carolina mantle slug mucus concentrations ranging from 1:0 to 1:100,000 v/v in ethanol (Figure 2C). Both genotypes exhibited similar concentration-dependent chemotaxis responses across the tested range, with no difference observed between wild-type and RB1526 animals. Taken together, these findings indicate that loss of srd-44 does not significantly alter chemotaxis toward the tested volatile odorants or Carolina mantle slug mucus under the conditions examined, providing an opportunity for further investigation of its sensory function.
","references":[{"reference":"Ballard KR, Klein AH, Hayes RA, Wang T, Cummins SF. 2021. The protein and volatile components of trail mucus in the Common Garden Snail, Cornu aspersum. PLoS One 16(5): e0251565.
","pubmedId":"34043643","doi":""},{"reference":"Bargmann CI, Hartwieg E, Horvitz HR. 1993. Odorant-selective genes and neurons mediate olfaction in C. elegans. Cell 74(3): 515-27.
","pubmedId":"8348618","doi":""},{"reference":"Frézal L, Félix MA. 2015. C. elegans outside the Petri dish. Elife 4: 10.7554/eLife.05849.
","pubmedId":"25822066","doi":""},{"reference":"Gaerlan M, Carrillo M, Ceva S, Chundi S, Diallo B, Fong JN, et al., O'Connell LA. 2025. Velvety tree ant extract is a chemotaxis repellent for C. elegans. MicroPubl Biol 2025: 10.17912/micropub.biology.001531.
","pubmedId":"40535527","doi":""},{"reference":"Jobe A, Vijayan R. 2024. Orphan G protein-coupled receptors: the ongoing search for a home. Front Pharmacol 15: 1349097.
","pubmedId":"38495099","doi":""},{"reference":"Kaplan JM, Horvitz HR. 1993. A dual mechanosensory and chemosensory neuron in Caenorhabditis elegans. Proc Natl Acad Sci U S A 90(6): 2227-31.
","pubmedId":"8460126","doi":""},{"reference":"Kim K, Sato K, Shibuya M, Zeiger DM, Butcher RA, Ragains JR, et al., Sengupta P. 2009. Two chemoreceptors mediate developmental effects of dauer pheromone in C. elegans. Science 326(5955): 994-8.
","pubmedId":"19797623","doi":""},{"reference":"Margie O, Palmer C, Chin-Sang I. 2013. C. elegans chemotaxis assay. J Vis Exp(74): e50069.
","pubmedId":"23644543","doi":""},{"reference":"Robertson HM, Thomas JH. 2006. The putative chemoreceptor families of C. elegans. WormBook: 1-12.
","pubmedId":"18050473","doi":""},{"reference":"Smith AM, Morin MC. 2002. Biochemical differences between trail mucus and adhesive mucus from marsh periwinkle snails. Biol Bull 203(3): 338-46.
","pubmedId":"12480724","doi":""},{"reference":"Sternberg PW, Van Auken K, Wang Q, Wright A, Yook K, Zarowiecki M, et al., Stein L. 2024. WormBase 2024: status and transitioning to Alliance infrastructure. Genetics 227(1): 10.1093/genetics/iyae050.
","pubmedId":"38573366","doi":""},{"reference":"Stiernagle T. 2006. Maintenance of C. elegans. WormBook: 1-11.
","pubmedId":"18050451","doi":""},{"reference":"The Alliance of Genome Resources Consortium, Aleksander SA, Anagnostopoulos AV, Antonazzo G, Arnaboldi V, Attrill H, et al., Zytkovicz. 2024. Updates to the Alliance of Genome Resources central infrastructure. GENETICS 227: 10.1093/genetics/iyae049.
","pubmedId":"","doi":"10.1093/genetics/iyae049"},{"reference":"Troemel ER, Chou JH, Dwyer ND, Colbert HA, Bargmann CI. 1995. Divergent seven transmembrane receptors are candidate chemosensory receptors in C. elegans. Cell 83(2): 207-18.
","pubmedId":"7585938","doi":""},{"reference":"Vidal B, Aghayeva U, Sun H, Wang C, Glenwinkel L, Bayer EA, Hobert O. 2018. An atlas of Caenorhabditis elegans chemoreceptor expression. PLoS Biol 16(1): e2004218.
","pubmedId":"29293491","doi":""},{"reference":"Wood J, Ferkey D. 2019. unc-42 regulates the expression of ASH terminal fate markers. MicroPubl Biol 2019: 10.17912/micropub.biology.000114.
","pubmedId":"32550468","doi":""}],"title":"Behavioral Screening of the Candidate Chemoreceptor Gene srd-44 Using Diacetyl and Carolina Mantle Slug Mucus in Caenorhabditis elegans
","reviews":[{"reviewer":{"displayName":"Erika Sorensen"},"openAcknowledgement":false,"status":{"submitted":true}}],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":null}]},{"id":"90050c36-a3f9-4d72-adb5-214e441befde","decision":"edit","abstract":"C. elegans detect volatile odorants through amphid sensory neurons that express diverse G protein-coupled receptors (GPCRs), many of which remain functionally uncharacterized. We evaluated the candidate chemosensory GPCR srd-44 using the RB1526 mutant strain in chemotaxis assays against diacetyl and Carolina mantle (Philomycus carolinianus) slug mucus. Wild-type and mutant animals exhibited similar chemotaxis responses to both stimuli across a range of concentrations. Under these conditions, these results do not support a role for srd-44 in chemotaxis toward diacetyl or Carolina mantle slug mucus. These findings contribute to the functional evaluation of an orphan chemosensory GPCR in C. elegans.
","acknowledgements":"We acknowledge wormbase (now Alliance of Genome Resources), the Caenorhabditis Genetics Center, and the C. elegans Gene Knockout Project at the Oklahoma Medical Research Foundation (part of the International C. elegans Gene Knockout Consortium) for their contributions that make research such as ours possible.
","authors":[{"affiliations":["Goucher College, Towson, MD, US"],"departments":["Biological Sciences"],"credit":["formalAnalysis","fundingAcquisition","resources","supervision","writing_originalDraft","visualization","writing_reviewEditing"],"email":"jay.garaycochea@goucher.edu","firstName":"Jay","lastName":"Garaycochea","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0009-0007-5724-2667"},{"affiliations":["Goucher College, Towson, MD, US"],"departments":["Biological Sciences"],"credit":["conceptualization","investigation","methodology"],"email":"milo.roth@Goucher.edu","firstName":"Milo","lastName":"Roth","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"Funding for the project was provided by the The Clara W. Claasen '25 Scholarship and start-up funds from Goucher College.
","image":{"url":"https://portal.micropublication.org/uploads/bb779c490f393b058b95d24f29e21c30.png"},"imageCaption":"A. Chemotaxis index (CI) of wild-type (N2) and RB1526 (srd-44(ok1831)) animals toward diacetyl (1:100 v/v in ethanol), Carolina mantle slug mucus extract (100 dilution), Isoamyl Alcohol (1:100 v/v in ethanol), 1-Octanol (1:10 v/v in ethanol), and ethanol control after 60 minutes. CI was calculated as (T − C) / (T + C), where T represents animals in the test quadrants and C represents animals in ethanol control quadrants. Data are shown as individual biological replicates with mean ± SD. Statistical analysis was performed using two-way ANOVA (genotype x stimulus) followed by Tukey's multiple comparisons test. B. CI of wild-type and RB1526 animals towards diacetyl concentrations ranging from 1.14 × 10⁻⁵ M to 1.14 M after 60 minutes. Data shown as mean ± SD from 3-10 biological replicates per concentration. C. CI of wild-type and RB1526 animals towards Carolina Mantle Slug mucus concentrations ranging from a 100 to 10-5 dilution from original sample after 60 minutes. Data shown as mean ± SD from 3-4 biological replicates per concentration. No consistent differences in chemotaxis responses were observed between wild-type and RB1526 animals across the concentration range tested.
","imageTitle":"Loss of srd-44 is not associated with reduced chemotaxis to various odorants
","methods":"N2 and RB1526 (srd-44(ok1831)) strains were provided by the CGC, which is funded by NIH Office of Research Infrastructure Programs (P40 OD010440). The srd-44(ok1831) allele contains a 2064 bp deletion withing the srd-44 (F17A2.8) locus on chromosome X and is predicted to cause transcript ablation of the primary srd-44 transcript (Alliance of Genome Resources Consortium, 2024; Sternberg et al., 2024). This allele was generated by the C. elegans Gene Knockout Project at the Oklahoma Medical Research Foundation, which was part of the International C. elegans Gene Knockout Consortium. The srd-44(ok1831) allele deletion was confirmed through a primer pair of inner left (cttgatcagtcgctctcgtg) and inner right (cgcaaccattttggagagac) following six backcrossings. C. elegans strains were maintained using standard nematode culture protocols as previously described (Stiernagle, 2006). Briefly, an OP50 E. coli lawn was grown on NGM-coated 60 mm petri plates. Picked eggs were transferred from freshly starved stock plates, incubated at 20°C for 2-3 days, then noted for presence of synchronized L4 and young adults. Worms were collected and pelleted using an M9 wash buffer before being transferred to a prepared testing plate.
Chemotaxis assay was adopted from Margie et al. (2013) by utilizing four quadrant divisions with two test and two control quadrants in a 10cm petri plate. 40-150 Pelleted worms were placed in the center and allowed to absorb into the agar surface for 5 minutes. Testing compounds, mixed with 0.5M sodium azide, were applied to designated areas. After one hour at room temperature, plates were placed into a 4°C cooler, then scored for worms that travelled 0.5 cm from the point of origin. Chemotaxis Index was calculated using the following equation: Chemotaxis Index = (# Worms in Both Test Quadrants - Worms in Both Control Quadrants) / (Total # of Scored Worms).
Philomycus carolinianus (Carolina Mantle slug) was collected from the deciduous forests surrounding Goucher College in Towson, Maryland USA. Slugs were placed in a terrarium for 24 hours before being transferred to a terrarium containing autoclaved dirt, rocks, and woody material. Slugs were fed a diet of autoclaved carrots and lettuce, misted with autoclaved tap water via a Honeywell humidifier, and allowed to acclimate for 3 days before mucus was collected. Cotton tipped applicators were used to collect mucus, then swirled in an Eppendorf tube with 45 µl of ethanol and stored in a 4°C cooler before being transferred to a -80°C freezer. Ethanol was selected as the solvent for mucus preparation as previous behavioral studies using chemically complex invertebrate samples have used ethanol and other solvents without detecting solvent dependent chemotactic responses (Archer et al., 2020). All slugs were returned to their native habitat at the end of the study.
Statistics were performed using Graphpad Prism software version 10.5.0 for windows. A two-way ANOVA was used to evaluate the effects of genotype and stimulus condition, followed by Tukey's multiple comparisons to assess genotype differences within each condition. Descriptive statistics were used to calculate mean and standard deviation. The figure was created using grouped summary data.
OpenAI's ChatGPT, version GPT-5.2, was used for editing language, clarity, and structure of the author's original draft manuscript of the description and abstract. Title, figures, figure legend, and methods were author originated. Prompts requesting deep research into previous experimental findings, suggestions for references, and critique of conclusions were manually reviewed and verified by the authors. The authors accept full responsibility for the accuracy of all content in the final manuscript.
","reagents":"Strain | Genotype | Available From |
CGC | ||
CGC |
Species | Common Name | Location Acquired |
Philomycus carolinianus | Carolina Mantle slug | Woods surrounding Goucher College |
Reagent | Source | Description |
1-Octanol, ≥99% (GC) | Sigma Aldrich | Cat# 472328-100ML |
Diacetyl, 98.0+% | TCI Chemicals | Cat# B0682-25ML |
Ethanol, 200 proof | Pharmco | Cat# 111000190-1Gal |
Isoamyl Alcohol, ≥98% (GC) | Sigma Aldrich | Cat# W205702-1KG-K |
Statistical Analysis
Tukey's multiple comparisons test (Wildtype vs. RB1526) | Predicted (LS) Mean diff. | Adjusted P Value |
Diacetyl | -0.2448 to 0.2867 | 0.8733 |
Carolina Mantleslug | -0.4895 to 0.1697 | 0.3300 |
Isoamyl Alcohol | -0.4793 to 0.06227 | 0.1265 |
1-Octanol | -0.07980 to 0.5368 | 0.1408 |
Ethanol | -0.3069 to 0.3522 | 0.8895 |
C. elegans detects volatile odorants through amphid sensory neurons that express diverse G protein-coupled receptors (GPCRs), many of which remain functionally uncharacterized(Bargmann et al., 1993; Robertson & Thomas, 2006). Members of this large chemosensory GPCR family mediate responses to both defined odorants and complex biological compounds(Bargmann et al., 1993; Kim et al., 2009a; Troemel et al., 1995a). Identifying the ligands of orphaned GPCRs is a challenging and time-consuming task that requires a comprehensive approach integrating a multitude of techniques to understand the full range of GPCR responses (Jobe & Vijayan, 2024). One potential approach is using biological complex compounds with gas chromatography/mass spectrometry to screen for candidate metabolites (Gaerlan et al., 2025).
C. elegans coexist in natural environments with terrestrial invertebrates including isopods and gastropods such as slugs (Frézal & Félix, 2015). Slug mucus contains a mixture of proteins, glycoproteins, and small metabolites that act as volatile compounds and may serve as multimodal chemical cues (Ballard et al., 2021; Smith & Morin, 2002). Members of the large chemosensory GPCR family expressed in amphid neurons mediate responses to both defined odorants and complex biological compounds (Bargmann et al., 1993; Kim et al., 2009b; Troemel et al., 1995b). The properties of slug mucus position it as a candidate for screening odorant GPCRs in C. elegans.
srd-44 encodes a predicted seven-transmembrane receptor belonging to the serpentine receptor class delta (srd) family, a group of genes enriched in chemosensory neurons(Robertson & Thomas, 2006; Troemel et al., 1995a; Vidal et al., 2018). A previous reporter-based study suggested expression of srd-44 in ASH amphid sensory neurons, although endogenous expression patterns remain incompletely defined (Wood & Ferkey, 2019). ASH neurons are canonically implicated in avoidance responses to nose touch and avoidance of noxious chemicals (Kaplan & Horvitz, 1993); however, the valence of chemosensory responses in C. elegans is not strictly fixed by neuron identity and can shift depending on stimulus concentration, context, and the specific receptor(s) engaged (Khan et al., 2022). Despite its predicted classification as a chemosensory receptor, no ligand or defined behavioral function has been reported for srd-44. Given its putative expression in ASH sensory neurons, srd-44 was selected as a candidate for behavioral screening to test its role in mediating responses to a panel of attractive and aversive chemosensory stimuli.
Wild-type (N2) and RB1526 (srd-44(ok1831))C. elegans were compared for chemotaxis towards the volatile attractant diacetyl and isoamyl alcohol, the volatile repellent 1-octanol, and the undefined cue Carolina mantle slug mucus, with ethanol as the negative control (Figure 1A). No significant differences in chemotaxis were observed between genotypes for any of the tested stimuli. To further evaluate responses to attractants, both strains were subsequently examined using diacetyl concentrations ranging from 1.14 × 10⁻⁵ M to 1.14 M (Figure 1B) and Carolina mantle slug mucus concentrations ranging from 1:0 to 1:100,000 v/v in ethanol (Figure 2C). Both genotypes exhibited similar concentration-dependent chemotaxis responses across the tested range, with no difference observed between wild-type and RB1526 animals. Taken together, these findings indicate that loss of srd-44 does not significantly alter chemotaxis toward the tested volatile odorants or Carolina mantle slug mucus under the conditions examined, providing an opportunity for further investigation of its sensory function.
","references":[{"reference":"Archer H, Deiparine S, Andersen EC. 2020. The nematode Caenorhabditis elegans and the terrestrial isopod Porcellio scaber likely interact opportunistically. PLoS One 15(6): e0235000.
","pubmedId":"32589676","doi":""},{"reference":"Ballard KR, Klein AH, Hayes RA, Wang T, Cummins SF. 2021. The protein and volatile components of trail mucus in the Common Garden Snail, Cornu aspersum. PLoS One 16(5): e0251565.
","pubmedId":"34043643","doi":""},{"reference":"Bargmann CI, Hartwieg E, Horvitz HR. 1993. Odorant-selective genes and neurons mediate olfaction in C. elegans. Cell 74(3): 515-27.
","pubmedId":"8348618","doi":""},{"reference":"Frézal L, Félix MA. 2015. C. elegans outside the Petri dish. Elife 4: 10.7554/eLife.05849.
","pubmedId":"25822066","doi":""},{"reference":"Gaerlan M, Carrillo M, Ceva S, Chundi S, Diallo B, Fong JN, et al., O'Connell LA. 2025. Velvety tree ant extract is a chemotaxis repellent for C. elegans. MicroPubl Biol 2025: 10.17912/micropub.biology.001531.
","pubmedId":"40535527","doi":""},{"reference":"Jobe A, Vijayan R. 2024. Orphan G protein-coupled receptors: the ongoing search for a home. Front Pharmacol 15: 1349097.
","pubmedId":"38495099","doi":""},{"reference":"Kaplan JM, Horvitz HR. 1993. A dual mechanosensory and chemosensory neuron in Caenorhabditis elegans. Proc Natl Acad Sci U S A 90(6): 2227-31.
","pubmedId":"8460126","doi":""},{"reference":"Khan M, Hartmann AH, O'Donnell MP, Piccione M, Pandey A, Chao PH, et al., Sengupta P. 2022. Context-dependent reversal of odorant preference is driven by inversion of the response in a single sensory neuron type. PLoS Biol 20(6): e3001677.
","pubmedId":"35696430","doi":""},{"reference":"Kim K, Sato K, Shibuya M, Zeiger DM, Butcher RA, Ragains JR, et al., Sengupta P. 2009. Two chemoreceptors mediate developmental effects of dauer pheromone in C. elegans. Science 326(5955): 994-8.
","pubmedId":"19797623","doi":""},{"reference":"Margie O, Palmer C, Chin-Sang I. 2013. C. elegans chemotaxis assay. J Vis Exp(74): e50069.
","pubmedId":"23644543","doi":""},{"reference":"Robertson HM, Thomas JH. 2006. The putative chemoreceptor families of C. elegans. WormBook: 1-12.
","pubmedId":"18050473","doi":""},{"reference":"Smith AM, Morin MC. 2002. Biochemical differences between trail mucus and adhesive mucus from marsh periwinkle snails. Biol Bull 203(3): 338-46.
","pubmedId":"12480724","doi":""},{"reference":"Sternberg PW, Van Auken K, Wang Q, Wright A, Yook K, Zarowiecki M, et al., Stein L. 2024. WormBase 2024: status and transitioning to Alliance infrastructure. Genetics 227(1): 10.1093/genetics/iyae050.
","pubmedId":"38573366","doi":""},{"reference":"Stiernagle T. 2006. Maintenance of C. elegans. WormBook: 1-11.
","pubmedId":"18050451","doi":""},{"reference":"The Alliance of Genome Resources Consortium, Aleksander SA, Anagnostopoulos AV, Antonazzo G, Arnaboldi V, Attrill H, et al., Zytkovicz. 2024. Updates to the Alliance of Genome Resources central infrastructure. GENETICS 227: 10.1093/genetics/iyae049.
","pubmedId":"","doi":"10.1093/genetics/iyae049"},{"reference":"Troemel ER, Chou JH, Dwyer ND, Colbert HA, Bargmann CI. 1995. Divergent seven transmembrane receptors are candidate chemosensory receptors in C. elegans. Cell 83(2): 207-18.
","pubmedId":"7585938","doi":""},{"reference":"Vidal B, Aghayeva U, Sun H, Wang C, Glenwinkel L, Bayer EA, Hobert O. 2018. An atlas of Caenorhabditis elegans chemoreceptor expression. PLoS Biol 16(1): e2004218.
","pubmedId":"29293491","doi":""},{"reference":"Wood J, Ferkey D. 2019. unc-42 regulates the expression of ASH terminal fate markers. MicroPubl Biol 2019: 10.17912/micropub.biology.000114.
","pubmedId":"32550468","doi":""}],"title":"Loss of the Orphan GPCR srd-44 Does Not Alter Chemotaxis of Caenorhabditis elegans to Carolina Mantle Slug Mucus
","reviews":[],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":"1790537086888"}]},{"id":"12454aa3-0dea-4a2f-b6da-4c63f67f0701","decision":"accept","abstract":"C. elegans detect volatile odorants through amphid sensory neurons that express diverse G protein-coupled receptors (GPCRs), many of which remain functionally uncharacterized. We evaluated the candidate chemosensory GPCR srd-44 using the RB1526 mutant strain in chemotaxis assays against diacetyl and Carolina mantle (Philomycus carolinianus) slug mucus. Wild-type and mutant animals exhibited similar chemotaxis responses to both stimuli across a range of concentrations. Under these conditions, these results do not support a role for srd-44 in chemotaxis toward diacetyl or Carolina mantle slug mucus. These findings contribute to the functional evaluation of an orphan chemosensory GPCR in C. elegans.
","acknowledgements":"We acknowledge wormbase (now Alliance of Genome Resources), the Caenorhabditis Genetics Center, and the C. elegans Gene Knockout Project at the Oklahoma Medical Research Foundation (part of the International C. elegans Gene Knockout Consortium) for their contributions that make research such as ours possible.
","authors":[{"affiliations":["Goucher College, Towson, MD, US"],"departments":["Biological Sciences"],"credit":["formalAnalysis","fundingAcquisition","resources","supervision","writing_originalDraft","visualization","writing_reviewEditing"],"email":"jay.garaycochea@goucher.edu","firstName":"Jay","lastName":"Garaycochea","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0009-0007-5724-2667"},{"affiliations":["Goucher College, Towson, MD, US"],"departments":["Biological Sciences"],"credit":["conceptualization","investigation","methodology"],"email":"milo.roth@Goucher.edu","firstName":"Milo","lastName":"Roth","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"Funding for the project was provided by the The Clara W. Claasen '25 Scholarship and start-up funds from Goucher College.
","image":{"url":"https://portal.micropublication.org/uploads/bb779c490f393b058b95d24f29e21c30.png"},"imageCaption":"A. Chemotaxis index (CI) of wild-type (N2) and RB1526 (srd-44(ok1831)) animals toward diacetyl (1:100 v/v in ethanol), Carolina mantle slug mucus extract (100 dilution), Isoamyl Alcohol (1:100 v/v in ethanol), 1-Octanol (1:10 v/v in ethanol), and ethanol control after 60 minutes. CI was calculated as (T − C) / (T + C), where T represents animals in the test quadrants and C represents animals in ethanol control quadrants. Data are shown as individual biological replicates with mean ± SD. Statistical analysis was performed using two-way ANOVA (genotype x stimulus) followed by Tukey's multiple comparisons test. B. CI of wild-type and RB1526 animals towards diacetyl concentrations ranging from 1.14 × 10⁻⁵ M to 1.14 M after 60 minutes. Data shown as mean ± SD from 3-10 biological replicates per concentration. C. CI of wild-type and RB1526 animals towards Carolina Mantle Slug mucus concentrations ranging from a 100 to 10-5 dilution from original sample after 60 minutes. Data shown as mean ± SD from 3-4 biological replicates per concentration. No consistent differences in chemotaxis responses were observed between wild-type and RB1526 animals across the concentration range tested.
","imageTitle":"Loss of srd-44 is not associated with reduced chemotaxis to various odorants
","methods":"N2 and RB1526 (srd-44(ok1831)) strains were provided by the CGC, which is funded by NIH Office of Research Infrastructure Programs (P40 OD010440). The srd-44(ok1831) allele contains a 2064 bp deletion withing the srd-44 (F17A2.8) locus on chromosome X and is predicted to cause transcript ablation of the primary srd-44 transcript (Alliance of Genome Resources Consortium, 2024; Sternberg et al., 2024). This allele was generated by the C. elegans Gene Knockout Project at the Oklahoma Medical Research Foundation, which was part of the International C. elegans Gene Knockout Consortium. The srd-44(ok1831) allele deletion was confirmed through a primer pair of inner left (cttgatcagtcgctctcgtg) and inner right (cgcaaccattttggagagac) following six backcrossings. C. elegans strains were maintained using standard nematode culture protocols as previously described (Stiernagle, 2006). Briefly, an OP50 E. coli lawn was grown on NGM-coated 60 mm petri plates. Picked eggs were transferred from freshly starved stock plates, incubated at 20°C for 2-3 days, then noted for presence of synchronized L4 and young adults. Worms were collected and pelleted using an M9 wash buffer before being transferred to a prepared testing plate.
Chemotaxis assay was adopted from Margie et al. (2013) by utilizing four quadrant divisions with two test and two control quadrants in a 10cm petri plate. 40-150 Pelleted worms were placed in the center and allowed to absorb into the agar surface for 5 minutes. Testing compounds, mixed with 0.5M sodium azide, were applied to designated areas. After one hour at room temperature, plates were placed into a 4°C cooler, then scored for worms that travelled 0.5 cm from the point of origin. Chemotaxis Index was calculated using the following equation: Chemotaxis Index = (# Worms in Both Test Quadrants - Worms in Both Control Quadrants) / (Total # of Scored Worms).
Philomycus carolinianus (Carolina Mantle slug) was collected from the deciduous forests surrounding Goucher College in Towson, Maryland USA. Slugs were placed in a terrarium for 24 hours before being transferred to a terrarium containing autoclaved dirt, rocks, and woody material. Slugs were fed a diet of autoclaved carrots and lettuce, misted with autoclaved tap water via a Honeywell humidifier, and allowed to acclimate for 3 days before mucus was collected. Cotton tipped applicators were used to collect mucus, then swirled in an Eppendorf tube with 45 µl of ethanol and stored in a 4°C cooler before being transferred to a -80°C freezer. Ethanol was selected as the solvent for mucus preparation as previous behavioral studies using chemically complex invertebrate samples have used ethanol and other solvents without detecting solvent dependent chemotactic responses (Archer et al., 2020). All slugs were returned to their native habitat at the end of the study.
Statistics were performed using Graphpad Prism software version 10.5.0 for windows. A two-way ANOVA was used to evaluate the effects of genotype and stimulus condition, followed by Tukey's multiple comparisons to assess genotype differences within each condition. Descriptive statistics were used to calculate mean and standard deviation. The figure was created using grouped summary data.
","reagents":"Strain | Genotype | Available From |
CGC | ||
CGC |
Species | Common Name | Location Acquired |
Philomycus carolinianus | Carolina Mantle slug | Woods surrounding Goucher College |
Reagent | Source | Description |
1-Octanol, ≥99% (GC) | Sigma Aldrich | Cat# 472328-100ML |
Diacetyl, 98.0+% | TCI Chemicals | Cat# B0682-25ML |
Ethanol, 200 proof | Pharmco | Cat# 111000190-1Gal |
Isoamyl Alcohol, ≥98% (GC) | Sigma Aldrich | Cat# W205702-1KG-K |
Statistical Analysis
Tukey's multiple comparisons test (Wildtype vs. RB1526) | Predicted (LS) Mean diff. | Adjusted P Value |
Diacetyl | -0.2448 to 0.2867 | 0.8733 |
Carolina Mantleslug | -0.4895 to 0.1697 | 0.3300 |
Isoamyl Alcohol | -0.4793 to 0.06227 | 0.1265 |
1-Octanol | -0.07980 to 0.5368 | 0.1408 |
Ethanol | -0.3069 to 0.3522 | 0.8895 |
C. elegans detects volatile odorants through amphid sensory neurons that express diverse G protein-coupled receptors (GPCRs), many of which remain functionally uncharacterized(Bargmann et al., 1993; Robertson & Thomas, 2006). Members of this large chemosensory GPCR family mediate responses to both defined odorants and complex biological compounds(Bargmann et al., 1993; Kim et al., 2009a; Troemel et al., 1995a). Identifying the ligands of orphaned GPCRs is a challenging and time-consuming task that requires a comprehensive approach integrating a multitude of techniques to understand the full range of GPCR responses (Jobe & Vijayan, 2024). One potential approach is using biological complex compounds with gas chromatography/mass spectrometry to screen for candidate metabolites (Gaerlan et al., 2025).
C. elegans coexist in natural environments with terrestrial invertebrates including isopods and gastropods such as slugs (Frézal & Félix, 2015). Slug mucus contains a mixture of proteins, glycoproteins, and small metabolites that act as volatile compounds and may serve as multimodal chemical cues (Ballard et al., 2021; Smith & Morin, 2002). Members of the large chemosensory GPCR family expressed in amphid neurons mediate responses to both defined odorants and complex biological compounds (Bargmann et al., 1993; Kim et al., 2009b; Troemel et al., 1995b). The properties of slug mucus position it as a candidate for screening odorant GPCRs in C. elegans.
srd-44 encodes a predicted seven-transmembrane receptor belonging to the serpentine receptor class delta (srd) family, a group of genes enriched in chemosensory neurons(Robertson & Thomas, 2006; Troemel et al., 1995a; Vidal et al., 2018). A previous reporter-based study suggested expression of srd-44 in ASH amphid sensory neurons, although endogenous expression patterns remain incompletely defined (Wood & Ferkey, 2019). ASH neurons are canonically implicated in avoidance responses to nose touch and avoidance of noxious chemicals (Kaplan & Horvitz, 1993); however, the valence of chemosensory responses in C. elegans is not strictly fixed by neuron identity and can shift depending on stimulus concentration, context, and the specific receptor(s) engaged (Khan et al., 2022). Despite its predicted classification as a chemosensory receptor, no ligand or defined behavioral function has been reported for srd-44. Given its putative expression in ASH sensory neurons, srd-44 was selected as a candidate for behavioral screening to test its role in mediating responses to a panel of attractive and aversive chemosensory stimuli.
Wild-type (N2) and RB1526 (srd-44(ok1831))C. elegans were compared for chemotaxis towards the volatile attractant diacetyl and isoamyl alcohol, the volatile repellent 1-octanol, and the undefined cue Carolina mantle slug mucus, with ethanol as the negative control (Figure 1A). No significant differences in chemotaxis were observed between genotypes for any of the tested stimuli. To further evaluate responses to attractants, both strains were subsequently examined using diacetyl concentrations ranging from 1.14 × 10⁻⁵ M to 1.14 M (Figure 1B) and Carolina mantle slug mucus concentrations ranging from 1:0 to 1:100,000 v/v in ethanol (Figure 2C). Both genotypes exhibited similar concentration-dependent chemotaxis responses across the tested range, with no difference observed between wild-type and RB1526 animals. Taken together, these findings indicate that loss of srd-44 does not significantly alter chemotaxis toward the tested volatile odorants or Carolina mantle slug mucus under the conditions examined, providing an opportunity for further investigation of its sensory function.
","references":[{"reference":"Archer H, Deiparine S, Andersen EC. 2020. The nematode Caenorhabditis elegans and the terrestrial isopod Porcellio scaber likely interact opportunistically. PLoS One 15(6): e0235000.
","pubmedId":"32589676","doi":""},{"reference":"Ballard KR, Klein AH, Hayes RA, Wang T, Cummins SF. 2021. The protein and volatile components of trail mucus in the Common Garden Snail, Cornu aspersum. PLoS One 16(5): e0251565.
","pubmedId":"34043643","doi":""},{"reference":"Bargmann CI, Hartwieg E, Horvitz HR. 1993. Odorant-selective genes and neurons mediate olfaction in C. elegans. Cell 74(3): 515-27.
","pubmedId":"8348618","doi":""},{"reference":"Frézal L, Félix MA. 2015. C. elegans outside the Petri dish. Elife 4: 10.7554/eLife.05849.
","pubmedId":"25822066","doi":""},{"reference":"Gaerlan M, Carrillo M, Ceva S, Chundi S, Diallo B, Fong JN, et al., O'Connell LA. 2025. Velvety tree ant extract is a chemotaxis repellent for C. elegans. MicroPubl Biol 2025: 10.17912/micropub.biology.001531.
","pubmedId":"40535527","doi":""},{"reference":"Jobe A, Vijayan R. 2024. Orphan G protein-coupled receptors: the ongoing search for a home. Front Pharmacol 15: 1349097.
","pubmedId":"38495099","doi":""},{"reference":"Kaplan JM, Horvitz HR. 1993. A dual mechanosensory and chemosensory neuron in Caenorhabditis elegans. Proc Natl Acad Sci U S A 90(6): 2227-31.
","pubmedId":"8460126","doi":""},{"reference":"Khan M, Hartmann AH, O'Donnell MP, Piccione M, Pandey A, Chao PH, et al., Sengupta P. 2022. Context-dependent reversal of odorant preference is driven by inversion of the response in a single sensory neuron type. PLoS Biol 20(6): e3001677.
","pubmedId":"35696430","doi":""},{"reference":"Kim K, Sato K, Shibuya M, Zeiger DM, Butcher RA, Ragains JR, et al., Sengupta P. 2009. Two chemoreceptors mediate developmental effects of dauer pheromone in C. elegans. Science 326(5955): 994-8.
","pubmedId":"19797623","doi":""},{"reference":"Margie O, Palmer C, Chin-Sang I. 2013. C. elegans chemotaxis assay. J Vis Exp(74): e50069.
","pubmedId":"23644543","doi":""},{"reference":"Robertson HM, Thomas JH. 2006. The putative chemoreceptor families of C. elegans. WormBook: 1-12.
","pubmedId":"18050473","doi":""},{"reference":"Smith AM, Morin MC. 2002. Biochemical differences between trail mucus and adhesive mucus from marsh periwinkle snails. Biol Bull 203(3): 338-46.
","pubmedId":"12480724","doi":""},{"reference":"Sternberg PW, Van Auken K, Wang Q, Wright A, Yook K, Zarowiecki M, et al., Stein L. 2024. WormBase 2024: status and transitioning to Alliance infrastructure. Genetics 227(1): 10.1093/genetics/iyae050.
","pubmedId":"38573366","doi":""},{"reference":"Stiernagle T. 2006. Maintenance of C. elegans. WormBook: 1-11.
","pubmedId":"18050451","doi":""},{"reference":"The Alliance of Genome Resources Consortium, Aleksander SA, Anagnostopoulos AV, Antonazzo G, Arnaboldi V, Attrill H, et al., Zytkovicz. 2024. Updates to the Alliance of Genome Resources central infrastructure. GENETICS 227: 10.1093/genetics/iyae049.
","pubmedId":"","doi":"10.1093/genetics/iyae049"},{"reference":"Troemel ER, Chou JH, Dwyer ND, Colbert HA, Bargmann CI. 1995. Divergent seven transmembrane receptors are candidate chemosensory receptors in C. elegans. Cell 83(2): 207-18.
","pubmedId":"7585938","doi":""},{"reference":"Vidal B, Aghayeva U, Sun H, Wang C, Glenwinkel L, Bayer EA, Hobert O. 2018. An atlas of Caenorhabditis elegans chemoreceptor expression. PLoS Biol 16(1): e2004218.
","pubmedId":"29293491","doi":""},{"reference":"Wood J, Ferkey D. 2019. unc-42 regulates the expression of ASH terminal fate markers. MicroPubl Biol 2019: 10.17912/micropub.biology.000114.
","pubmedId":"32550468","doi":""}],"title":"Loss of the Orphan GPCR srd-44 Does Not Alter Chemotaxis of Caenorhabditis elegans to Carolina Mantle Slug Mucus
","reviews":[],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":null}]},{"id":"59468d6b-bd4b-41d4-8256-9392bd70764f","decision":"publish","abstract":"C. elegans detect volatile odorants through amphid sensory neurons that express diverse G protein-coupled receptors (GPCRs), many of which remain functionally uncharacterized. We evaluated the candidate chemosensory GPCR srd-44 using the RB1526 mutant strain in chemotaxis assays against diacetyl and Carolina mantle (Philomycus carolinianus) slug mucus. Wild-type and mutant animals exhibited similar chemotaxis responses to both stimuli across a range of concentrations. Under these conditions, these results do not support a role for srd-44 in chemotaxis toward diacetyl or Carolina mantle slug mucus. These findings contribute to the functional evaluation of an orphan chemosensory GPCR in C. elegans.
","acknowledgements":"We acknowledge wormbase (now Alliance of Genome Resources), the Caenorhabditis Genetics Center, and the C. elegans Gene Knockout Project at the Oklahoma Medical Research Foundation (part of the International C. elegans Gene Knockout Consortium) for their contributions that make research such as ours possible.
","authors":[{"affiliations":["Goucher College, Towson, MD, US"],"departments":["Biological Sciences"],"credit":["formalAnalysis","fundingAcquisition","resources","supervision","writing_originalDraft","visualization","writing_reviewEditing"],"email":"jay.garaycochea@goucher.edu","firstName":"Jay","lastName":"Garaycochea","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"0009-0007-5724-2667"},{"affiliations":["Goucher College, Towson, MD, US"],"departments":["Biological Sciences"],"credit":["conceptualization","investigation","methodology"],"email":"milo.roth@Goucher.edu","firstName":"Milo","lastName":"Roth","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"Funding for the project was provided by the The Clara W. Claasen '25 Scholarship and start-up funds from Goucher College.
","image":{"url":"https://portal.micropublication.org/uploads/bb779c490f393b058b95d24f29e21c30.png"},"imageCaption":"A. Chemotaxis index (CI) of wild-type (N2) and RB1526 (srd-44(ok1831)) animals toward diacetyl (1:100 v/v in ethanol), Carolina mantle slug mucus extract (100 dilution), Isoamyl Alcohol (1:100 v/v in ethanol), 1-Octanol (1:10 v/v in ethanol), and ethanol control after 60 minutes. CI was calculated as (T − C) / (T + C), where T represents animals in the test quadrants and C represents animals in ethanol control quadrants. Data are shown as individual biological replicates with mean ± SD. Statistical analysis was performed using two-way ANOVA (genotype x stimulus) followed by Tukey's multiple comparisons test. B. CI of wild-type and RB1526 animals towards diacetyl concentrations ranging from 1.14 × 10⁻⁵ M to 1.14 M after 60 minutes. Data shown as mean ± SD from 3-10 biological replicates per concentration. C. CI of wild-type and RB1526 animals towards Carolina Mantle Slug mucus concentrations ranging from a 100 to 10-5 dilution from original sample after 60 minutes. Data shown as mean ± SD from 3-4 biological replicates per concentration. No consistent differences in chemotaxis responses were observed between wild-type and RB1526 animals across the concentration range tested.
","imageTitle":"Loss of srd-44 is not associated with reduced chemotaxis to various odorants
","methods":"N2 and RB1526 (srd-44(ok1831)) strains were provided by the CGC, which is funded by NIH Office of Research Infrastructure Programs (P40 OD010440). The srd-44(ok1831) allele contains a 2064 bp deletion withing the srd-44 (F17A2.8) locus on chromosome X and is predicted to cause transcript ablation of the primary srd-44 transcript (Alliance of Genome Resources Consortium, 2024; Sternberg et al., 2024). This allele was generated by the C. elegans Gene Knockout Project at the Oklahoma Medical Research Foundation, which was part of the International C. elegans Gene Knockout Consortium. The srd-44(ok1831) allele deletion was confirmed through a primer pair of inner left (cttgatcagtcgctctcgtg) and inner right (cgcaaccattttggagagac) following six backcrossings. C. elegans strains were maintained using standard nematode culture protocols as previously described (Stiernagle, 2006). Briefly, an OP50 E. coli lawn was grown on NGM-coated 60 mm petri plates. Picked eggs were transferred from freshly starved stock plates, incubated at 20°C for 2-3 days, then noted for presence of synchronized L4 and young adults. Worms were collected and pelleted using an M9 wash buffer before being transferred to a prepared testing plate.
Chemotaxis assay was adopted from Margie et al. (2013) by utilizing four quadrant divisions with two test and two control quadrants in a 10cm petri plate. 40-150 Pelleted worms were placed in the center and allowed to absorb into the agar surface for 5 minutes. Testing compounds, mixed with 0.5M sodium azide, were applied to designated areas. After one hour at room temperature, plates were placed into a 4°C cooler, then scored for worms that travelled 0.5 cm from the point of origin. Chemotaxis Index was calculated using the following equation: Chemotaxis Index = (# Worms in Both Test Quadrants - Worms in Both Control Quadrants) / (Total # of Scored Worms).
Philomycus carolinianus (Carolina Mantle slug) was collected from the deciduous forests surrounding Goucher College in Towson, Maryland USA. Slugs were placed in a terrarium for 24 hours before being transferred to a terrarium containing autoclaved dirt, rocks, and woody material. Slugs were fed a diet of autoclaved carrots and lettuce, misted with autoclaved tap water via a Honeywell humidifier, and allowed to acclimate for 3 days before mucus was collected. Cotton tipped applicators were used to collect mucus, then swirled in an Eppendorf tube with 45 µl of ethanol and stored in a 4°C cooler before being transferred to a -80°C freezer. Ethanol was selected as the solvent for mucus preparation as previous behavioral studies using chemically complex invertebrate samples have used ethanol and other solvents without detecting solvent dependent chemotactic responses (Archer et al., 2020). All slugs were returned to their native habitat at the end of the study.
Statistics were performed using Graphpad Prism software version 10.5.0 for windows. A two-way ANOVA was used to evaluate the effects of genotype and stimulus condition, followed by Tukey's multiple comparisons to assess genotype differences within each condition. Descriptive statistics were used to calculate mean and standard deviation. The figure was created using grouped summary data.
","reagents":"Strain | Genotype | Available From |
CGC | ||
CGC |
Species | Common Name | Location Acquired |
Philomycus carolinianus | Carolina Mantle slug | Woods surrounding Goucher College |
Reagent | Source | Description |
1-Octanol, ≥99% (GC) | Sigma Aldrich | Cat# 472328-100ML |
Diacetyl, 98.0+% | TCI Chemicals | Cat# B0682-25ML |
Ethanol, 200 proof | Pharmco | Cat# 111000190-1Gal |
Isoamyl Alcohol, ≥98% (GC) | Sigma Aldrich | Cat# W205702-1KG-K |
Statistical Analysis
Tukey's multiple comparisons test (Wildtype vs. RB1526) | Predicted (LS) Mean diff. | Adjusted P Value |
Diacetyl | -0.2448 to 0.2867 | 0.8733 |
Carolina Mantleslug | -0.4895 to 0.1697 | 0.3300 |
Isoamyl Alcohol | -0.4793 to 0.06227 | 0.1265 |
1-Octanol | -0.07980 to 0.5368 | 0.1408 |
Ethanol | -0.3069 to 0.3522 | 0.8895 |
C. elegans detects volatile odorants through amphid sensory neurons that express diverse G protein-coupled receptors (GPCRs), many of which remain functionally uncharacterized (Bargmann et al., 1993; Robertson & Thomas, 2006). Members of this large chemosensory GPCR family mediate responses to both defined odorants and complex biological compounds (Bargmann et al., 1993; Kim et al., 2009a; Troemel et al., 1995a). Identifying the ligands of orphaned GPCRs is a challenging and time-consuming task that requires a comprehensive approach integrating a multitude of techniques to understand the full range of GPCR responses (Jobe & Vijayan, 2024). One potential approach is using biological complex compounds with gas chromatography/mass spectrometry to screen for candidate metabolites (Gaerlan et al., 2025).
C. elegans coexist in natural environments with terrestrial invertebrates including isopods and gastropods such as slugs (Frézal & Félix, 2015). Slug mucus contains a mixture of proteins, glycoproteins, and small metabolites that act as volatile compounds and may serve as multimodal chemical cues (Ballard et al., 2021; Smith & Morin, 2002). Members of the large chemosensory GPCR family expressed in amphid neurons mediate responses to both defined odorants and complex biological compounds (Bargmann et al., 1993; Kim et al., 2009b; Troemel et al., 1995b). The properties of slug mucus position it as a candidate for screening odorant GPCRs in C. elegans.
srd-44 encodes a predicted seven-transmembrane receptor belonging to the serpentine receptor class delta (srd) family, a group of genes enriched in chemosensory neurons (Robertson & Thomas, 2006; Troemel et al., 1995a; Vidal et al., 2018). A previous reporter-based study suggested expression of srd-44 in ASH amphid sensory neurons, although endogenous expression patterns remain incompletely defined (Wood & Ferkey, 2019). ASH neurons are canonically implicated in avoidance responses to nose touch and avoidance of noxious chemicals (Kaplan & Horvitz, 1993); however, the valence of chemosensory responses in C. elegans is not strictly fixed by neuron identity and can shift depending on stimulus concentration, context, and the specific receptor(s) engaged (Khan et al., 2022). Despite its predicted classification as a chemosensory receptor, no ligand or defined behavioral function has been reported for srd-44. Given its putative expression in ASH sensory neurons, srd-44 was selected as a candidate for behavioral screening to test its role in mediating responses to a panel of attractive and aversive chemosensory stimuli.
Wild-type (N2) and RB1526 (srd-44(ok1831)) C. elegans were compared for chemotaxis towards the volatile attractant diacetyl and isoamyl alcohol, the volatile repellent 1-octanol, and the undefined cue Carolina mantle slug mucus, with ethanol as the negative control (Figure 1A). No significant differences in chemotaxis were observed between genotypes for any of the tested stimuli. To further evaluate responses to attractants, both strains were subsequently examined using diacetyl concentrations ranging from 1.14 × 10⁻⁵ M to 1.14 M (Figure 1B) and Carolina mantle slug mucus concentrations ranging from 1:0 to 1:100,000 v/v in ethanol (Figure 2C). Both genotypes exhibited similar concentration-dependent chemotaxis responses across the tested range, with no difference observed between wild-type and RB1526 animals. Taken together, these findings indicate that loss of srd-44 does not significantly alter chemotaxis toward the tested volatile odorants or Carolina mantle slug mucus under the conditions examined, providing an opportunity for further investigation of its sensory function.
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","pubmedId":"32550468","doi":""}],"title":"Loss of the Orphan GPCR srd-44 Does Not Alter Chemotaxis of Caenorhabditis elegans to Carolina Mantle Slug Mucus
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