Sex-specific differences can influence aging and disease onset. Using the simple model organism Caenorhabditis elegans, we previously found that hermaphroditic nematodes can activate a robust mitochondrial unfolded protein response (UPRmt) in the intestines, while female nematodes cannot. We hypothesized that the accumulation of vitellogenins in the intestines of female nematodes may be suppressing the UPRmt. Loss of vitellogenins via RNAi enhanced the UPRmt in hermaphrodites but had no impact on the UPRmt of females unless a combination of vitellogenins was simultaneously knocked down. Furthermore, combinatorial loss of vitellogenins did not restore the female UPRmt to hermaphroditic levels. We conclude that additional factors besides vitellogenin content are involved in suppressing the intestinal UPRmt in female nematodes.
","acknowledgements":"We thank the Caenorhabditis Genetics Center (CGC), which is funded by the NIH Office of Research Infrastructure Programs (P40 OD010440).
","authors":[{"affiliations":["University of Maine"],"departments":["Molecular and Biomedical Sciences"],"credit":["dataCuration","methodology","validation","writing_originalDraft","formalAnalysis"],"email":"nathaniel.a.jordan@maine.edu","firstName":"Nathanial A.","lastName":"Jordan","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null},{"affiliations":["University of Maine"],"departments":["Molecular and Biomedical Sciences"],"credit":["conceptualization","fundingAcquisition","supervision","writing_reviewEditing"],"email":"suzanne.angeli@maine.edu","firstName":"Suzanne","lastName":"Angeli","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[{"awardId":"5P20GM144265 - 03 ","funderName":"National Institute of General Medical Sciences (United States)","awardRecipient":"Suzanne Angeli"}],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"NIGMS COBRE Research Pilot Project Subaward 5P20GM144265 - 03 and the UMaine Institute of Medicine.
","image":{"url":"https://portal.micropublication.org/uploads/d6b2a6ea0e651800efdd64682967f9f4.png"},"imageCaption":"A-D. Quantification of GFP intensity from phsp-6::GFP or fem-1(hc17); phsp-6::GFP nematodes. For single vitellogenin gene knockdown, nematodes were developed on either control vector (CV) RNAi or vit-1 (A), vit-3 (B), or vit-5 (C) RNAi at 25°C. At the young adult stage, control worms were shifted to 2o°C and onto RNAi plates seeded with CV or CV/atp-3 (50%/50% RNAi mixture). Vitellogenin RNAi treated worms were shifted onto RNAi plates seeded with the indicated CV/vit RNAi or vit/atp-3 (50%/50% RNAi mixture). For the combinatorial vitellogenin gene knockdown (D), nematodes were developed on either control vector (CV) RNAi or vit-1, vit-3, vit-5 (33%/33%/33% mixture) RNAi at 25°C. At the young adult stage, vitellogenin RNAi treated worms were shifted to 2o°C and onto RNAi plates seeded with the CV/vit-1,3,5 RNAi (50%/50% combinatorial RNAi mixture) or vit-1,3,5/atp-3 (50% combinatorial mixture/50% atp-3 RNAi). Error bars represent ± SEM. ***p ≤ 0.001, **p≤ 0.01, and *p≤ 0.05.
","imageTitle":"Impact of vitellogenin RNAi on the UPRmt
","methods":"Strains. Bristol N2 (wild type) nematodes were obtained from the Caenorhabditis Genetics Center (CGC, University of Minnesota) and cultured using standard conditions (Sulston J 1988). The following strains were used: GL347 (SJ4100 phsp-6::GFP backcrossed 6× to N2) and GL364 (fem-1(hc17); phsp-6::GFP).
Nematode and bacterial culture conditions. Nematodes were maintained on nematode growth medium (NGM) plates. NGM plates were seeded with Escherichia coli OP50 obtained from CGC that was grown in LB at 37°C for 18 hours shaking at 225 rpm. Seeded plates were dried for 48 hours at room temperature before use. For RNAi experiments, HT115 (DE3) bacteria obtained from the Horizon Biosciences RNAi library were used. All RNAi clones were verified via sequencing. RNAi plates were prepared by cooling NGM to 55°C and supplementing with a final concentration of 50μg/ml carbenicillin and 1 mM isopropyl β-d-1-thiogalactopyranoside (IPTG). RNAi bacteria were inoculated with one colony of RNAi bacteria into LB with 50μg/ml carbenicillin and were grown shaking overnight for 18 hours at 37° at 225 rpm. RNAi cultures were seeded on RNAi plates and allowed to grow for 48 hours at room temperature. Plates were stored at 4 °C for no longer than 2 weeks.
Microscopy. Nematodes were anesthetized with 2 -5mM levamisole and mounted on 2% agarose pads on glass slides. Fluorescence micrographs of GFP were taken using a Zeiss Axioscope 5 fluorescent compound microscope equipped with Zen microscopy imaging program. GFP expression was enhanced using the brightness/contrast tool in Adobe Photoshop. The same parameters were used for all images. GFP intensity of nematodes was quantified using ImageJ 1.54G. The “integrated density” of GFP expression and length of nematodes was measured using ImageJ tools. Integrated density value was normalized by number of nematodes and average length of nematodes. The final value is in arbitrary units.
Statistics. Significance between control and experimental groups was determined by using a two-tailed Student’s t-test. Asterisks denote corresponding statistical significance: *p < 0.05; **p < 0.01; ***p < 0.001. Error bars were generated using the standard error of the mean (SEM), typically from three or more pooled biological replicates.
","reagents":"","patternDescription":"Age-related diseases, such as cardiovascular disease, cancer, and neurodegenerative syndromes can display sex-specific differences. Since mitochondrial dysfunction is central to most age-related diseases, we examine the mitochondrial unfolded protein response (UPRmt), a conserved, broad-range transcriptional response that, among other functions, aids in the refolding of mitochondrial matrix proteins (Kim, Ramalho, and Haynes 2024). In C. elegans, UPRmt activation is largely localized to the intestine, the major metabolic tissue in the nematode. In addition to digestion, the intestine also carries out many functions similar to the liver, such as detoxification, immunity, and fat metabolism (Dimov and Maduro 2019). We and others recently discovered that the activation of the intestinal UPRmt depends on the function of another tissue: the germline (Foulger et al. 2025; Charmpilas et al. 2024; Shen et al. 2024; Zhou et al. 2024). Specifically, we found that adult nematodes with actively proliferating germlines, such as hermaphrodites or mated females, can activate a robust UPRmt in the intestines when challenged with either a high dose of the metal manganese (Mn) or RNAi of the OSCP/atp-3 subunit of complex V (Foulger et al. 2025). Conversely, adult nematodes with mutations that lead to a lack germline stem cells or lack of sperm, such glp-1 or fem-1 mutants, cannot activate the intestinal UPRmt. Additionally, we found that loss of the transcription factor FOXO/daf-16 fully restored the UPRmt in fem-1 mutants (Foulger et al. 2025). Here, we probe whether vitellogenesis, which occurs in the intestines and is dependent on FOXO/daf-16, impacts the UPRmt.
Vitellogenins are precursor yolk proteins that provide nutrients to developing embryos. In C. elegans, the highly conserved genes vit-1 through vit-6 are responsible for encoding these proteins (Perez and Lehner 2019). In hermaphrodites, they are synthesized in the intestines and transported into the pseudocoelom and to maturing oocytes (Perez and Lehner 2019). Young adult fem-1 mutants display elevated yolk content compared to wild-type hermaphrodites at Day 1 of adulthood (DePina et al. 2011), presumably due to a lack of vitellogenin transport out of the intestines to maturing oocytes. We used RNAi to knockdown vit-1, vit-3, vit-5, or all three genes simultaneously (vit-1,3,5) and monitored intestinal activation of the UPRmt in young adult nematodes. We used the UPRmt reporter in which the GFP expression is driven by the promoter of the mitochondrial chaperone mtHsp70/hsp-6 (Yoneda et al. 2004). We additionally used RNAi of OSCP/atp-3 to induce the UPRmt in young adult nematodes as previously described (Angeli et al. 2021). We found that in hermaphroditic nematodes, knockdown of vit-1 trended toward elevating the intestinal UPRmt (p=0.0504) and knockdown of vit-3, vit-5, or vit-1,3,5 all significantly elevated the UPRmt (Figure 1A-1D). In contrast, knockdown of vit-1, vit-3, or vit-5 did not significantly alter the UPRmt of fem-1 mutants (Figure 1A-1D). The knockdown of vit-1,3,5 did significantly enhance the UPRmt of the fem-1 mutants, but not to wild-types levels (Figure 1D). One caveat of this study is that we were not able to procure RNAi clones for vit-2, vit-4, or vit-6. However, since vit-2 is over 80% identical to vit-1 and vit-3/vit-4 are duplicated genes that are 99% identical (Perez and Lehner 2019), we would expect to observe more impact from individual RNAi knockdown on females if these vitellogenins played an outsized role on the UPRmt. The vit-6 gene is the most divergent vitellogenin, so it remains possible that it plays an untested role in the UPRmt. Overall, while it does appear that vitellogenins can mildly suppress the adult UPRmt, especially in hermaphroditic nematodes, we conclude that additional germline-to-intestinal signals are responsible for the potent suppression of the UPRmt in female nematodes.
","references":[{"reference":"Angeli S, Foulger A, Chamoli M, Peiris TH, Gerencser A, Shahmirzadi AA, Andersen J, Lithgow G. 2021. The mitochondrial permeability transition pore activates the mitochondrial unfolded protein response and promotes aging. eLife 10: 10.7554/elife.63453.
","pubmedId":"","doi":"10.7554/eLife.63453"},{"reference":"Charmpilas N, Sotiriou A, Axarlis K, Tavernarakis N, Hoppe T. 2024. Reproductive regulation of the mitochondrial stress response in Caenorhabditis elegans. Cell Reports 43: 114336.
","pubmedId":"","doi":"10.1016/j.celrep.2024.114336"},{"reference":"DePina AS, Iser WB, Park SS, Maudsley S, Wilson MA, Wolkow CA. 2011. Regulation of Caenorhabditis elegans vitellogenesis by DAF-2/IIS through separable transcriptional and posttranscriptional mechanisms. BMC Physiology 11: 10.1186/1472-6793-11-11.
","pubmedId":"","doi":"10.1186/1472-6793-11-11"},{"reference":"Dimov I, Maduro MF. 2019. The C. elegans intestine: organogenesis, digestion, and physiology. Cell and Tissue Research 377: 383-396.
","pubmedId":"","doi":" 10.1007/s00441-019-03036-4"},{"reference":"Foulger AC, Jordan NA, Castle A, Bhaumik D, Andersen JK, Lithgow GJ, Angeli S. 2025. Germline regulation of the intestinal mitochondrial unfolded protein response. GeroScience : 10.1007/s11357-025-01890-5.
","pubmedId":"","doi":"10.1007/s11357-025-01890-5"},{"reference":"Kim S, Ramalho TR, Haynes CM. 2024. Regulation of proteostasis and innate immunity via mitochondria-nuclear communication. Journal of Cell Biology 223: 10.1083/jcb.202310005.
","pubmedId":"","doi":"10.1083/jcb.202310005"},{"reference":"Perez MF, Lehner B. 2019. Vitellogenins - Yolk Gene Function and Regulation in Caenorhabditis elegans. Frontiers in Physiology 10: 10.3389/fphys.2019.01067.
","pubmedId":"","doi":"10.3389/fphys.2019.01067"},{"reference":"Shen K, Durieux J, Mena CG, Webster BM, Tsui CK, Zhang H, et al., Dillin. 2024. The germline coordinates mitokine signaling. Cell 187: 4605-4620.e17.
","pubmedId":"","doi":" 10.1016/j.cell.2024.06.010"},{"reference":"Sulston J, H.J., Methods, in The Nematode Caenorhabditis elegans, W.B. W., Editor. 1988, Cold Spring Harbor Laboratory Press Cold Spring Harbor. p. 587-606
","pubmedId":"","doi":""},{"reference":"Yoneda T, Benedetti C, Urano F, Clark SG, Harding HP, Ron D. 2004. Compartment-specific perturbation of protein handling activates genes encoding mitochondrial chaperones. Journal of Cell Science 117: 4055-4066.
","pubmedId":"","doi":" 10.1242/jcs.01275"},{"reference":"Zhou L, Jiang L, Li L, Ma C, Xia P, Ding W, Liu Y. 2024. A germline-to-soma signal triggers an age-related decline of mitochondrial stress response. Nature Communications 15: 10.1038/s41467-024-53064-0.
","pubmedId":"","doi":" 10.1038/s41467-024-53064-0"}],"title":"Role of vitellogenins on the mitochondrial unfolded protein response
","reviews":[{"reviewer":{"displayName":"Mark Pellegrino"},"openAcknowledgement":false,"status":{"submitted":true}}],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":null}]},{"id":"622bf716-597e-4d86-b109-341d96dea97f","decision":"accept","abstract":"Sex-specific differences influence aging and disease onset. Using Caenorhabditis elegans, we previously found that hermaphroditic nematodes activate a robust mitochondrial unfolded protein response (UPRmt) in the intestines, while female nematodes do not. We hypothesized that the accumulation of vitellogenins in females was suppressing the UPRmt. Loss of vitellogenins via RNAi enhanced the UPRmt in hermaphrodites but had no impact on the UPRmt of females unless a combination of vitellogenins was simultaneously knocked down. Furthermore, combinatorial loss of vitellogenins did not restore the female UPRmt to hermaphroditic levels. We conclude that additional factors besides vitellogenins are involved in suppressing the intestinal UPRmt in female nematodes.
","acknowledgements":"We thank the Caenorhabditis Genetics Center (CGC), which is funded by the NIH Office of Research Infrastructure Programs (P40 OD010440).
","authors":[{"affiliations":["University of Maine"],"departments":["Molecular and Biomedical Sciences"],"credit":["dataCuration","methodology","validation","writing_originalDraft","formalAnalysis"],"email":"nathaniel.a.jordan@maine.edu","firstName":"Nathanial A.","lastName":"Jordan","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null},{"affiliations":["University of Maine"],"departments":["Molecular and Biomedical Sciences"],"credit":["conceptualization","fundingAcquisition","supervision","writing_reviewEditing"],"email":"suzanne.angeli@maine.edu","firstName":"Suzanne","lastName":"Angeli","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[{"awardId":"5P20GM144265 - 03 ","funderName":"National Institute of General Medical Sciences (United States)","awardRecipient":"Suzanne Angeli"}],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"NIGMS COBRE Research Pilot Project Subaward 5P20GM144265 - 03 and the UMaine Institute of Medicine.
","image":{"url":"https://portal.micropublication.org/uploads/d6b2a6ea0e651800efdd64682967f9f4.png"},"imageCaption":"A-D. Quantification of GFP intensity from phsp-6::GFP or fem-1(hc17); phsp-6::GFP nematodes. For single vitellogenin gene knockdown, nematodes were developed on either control vector (CV) RNAi or vit-1 (A), vit-3 (B), or vit-5 (C) RNAi at 25°C. At the young adult stage, control worms were shifted to 2o°C and onto RNAi plates seeded with CV or CV/atp-3 (50%/50% RNAi mixture). Vitellogenin RNAi treated worms were shifted to 2o°C and onto RNAi plates seeded with the indicated CV/vit RNAi or vit/atp-3 (50%/50% RNAi mixture). For the combinatorial vitellogenin gene knockdown (D), nematodes were developed on either control vector (CV) RNAi or vit-1, vit-3, vit-5 (33%/33%/33% mixture) RNAi at 25°C. At the young adult stage, vitellogenin RNAi treated worms were shifted to 2o°C and onto RNAi plates seeded with the CV/vit-1,3,5 RNAi (50%/50% combinatorial RNAi mixture) or vit-1,3,5/atp-3 (50% combinatorial mixture/50% atp-3 RNAi). Error bars represent ± SEM. ***p ≤ 0.001, **p≤ 0.01, and *p≤ 0.05.
","imageTitle":"Impact of vitellogenin RNAi on the UPRmt
","methods":"Strains. Bristol N2 (wild type) nematodes were obtained from the Caenorhabditis Genetics Center (CGC, University of Minnesota) and cultured using standard conditions (Sulston J 1988). The following strains were used: GL347 (SJ4100 phsp-6::GFP backcrossed 6× to N2) and GL364 (fem-1(hc17); phsp-6::GFP).
Nematode and bacterial culture conditions. Nematodes were maintained on nematode growth medium (NGM) plates. NGM plates were seeded with Escherichia coli OP50 obtained from CGC that was grown in LB at 37°C for 18 hours shaking at 225 rpm. Seeded plates were dried for 48 hours at room temperature before use. For RNAi experiments, HT115 (DE3) bacteria obtained from the Horizon Biosciences RNAi library were used. All RNAi clones were verified via sequencing. RNAi plates were prepared by cooling NGM to 55°C and supplementing with a final concentration of 50μg/ml carbenicillin and 1 mM isopropyl β-d-1-thiogalactopyranoside (IPTG). RNAi bacteria were inoculated with one colony of RNAi bacteria into LB with 50μg/ml carbenicillin and were grown shaking overnight for 18 hours at 37° at 225 rpm. RNAi cultures were seeded on RNAi plates and allowed to grow for 48 hours at room temperature. Plates were stored at 4 °C for no longer than 2 weeks.
Microscopy. Nematodes were anesthetized with 2 - 5mM levamisole and mounted on 2% agarose pads on glass slides. Fluorescence micrographs of GFP were taken using a Zeiss Axioscope 5 fluorescent compound microscope equipped with Zen microscopy imaging program. GFP expression was enhanced using the brightness/contrast tool in Adobe Photoshop. The same parameters were used for all images. GFP intensity of nematodes was quantified using ImageJ 1.54G. The “integrated density” of GFP expression and length of nematodes was measured using ImageJ tools. Integrated density value was normalized by number of nematodes and average length of nematodes. The final value is in arbitrary units.
Statistics. Significance between control and experimental groups was determined by using a two-tailed Student's t-test. Asterisks denote corresponding statistical significance: *p < 0.05; **p < 0.01; ***p < 0.001. Error bars were generated using the standard error of the mean (SEM), typically from three or more pooled biological replicates.
","reagents":"","patternDescription":"Age-related diseases, such as cardiovascular disease, cancer, and neurodegenerative syndromes can display sex-specific differences. Since mitochondrial dysfunction is central to most age-related diseases, we examine the mitochondrial unfolded protein response (UPRmt), a conserved, broad-range transcriptional response that, among other functions, aids in the refolding of mitochondrial matrix proteins (Kim, Ramalho, and Haynes 2024). In C. elegans, UPRmt activation is largely localized to the intestine, the major metabolic tissue in the nematode. In addition to digestion, the intestine also carries out many functions similar to the liver, such as detoxification, immunity, and fat metabolism (Dimov and Maduro 2019). We and others recently discovered that the activation of the intestinal UPRmt depends on the function of another tissue: the germline (Foulger et al. 2025; Charmpilas et al. 2024; Shen et al. 2024; Zhou et al. 2024). Specifically, we found that adult nematodes with actively proliferating germlines, such as hermaphrodites or mated females, can activate a robust UPRmt in the intestines when challenged with either a high dose of the metal manganese (Mn) or RNAi of the OSCP/atp-3 subunit of complex V (Foulger et al. 2025). Conversely, adult nematodes with mutations that lead to a lack germline stem cells or lack of sperm, such glp-1 or fem-1 mutants, cannot activate the intestinal UPRmt. Additionally, we found that loss of the transcription factor FOXO/daf-16 fully restored the UPRmt in fem-1 mutants (Foulger et al. 2025). Here, we probe whether vitellogenesis, which occurs in the intestines and is dependent on FOXO/daf-16, impacts the UPRmt.
Vitellogenins are precursor yolk proteins that provide nutrients to developing embryos. In C. elegans, the highly conserved genes vit-1 through vit-6 are responsible for encoding these proteins (Perez and Lehner 2019). In hermaphrodites, they are synthesized in the intestines and transported into the pseudocoelom and to maturing oocytes (Perez and Lehner 2019). Young adult fem-1 mutants display elevated yolk content compared to wild-type hermaphrodites at Day 1 of adulthood (DePina et al. 2011), presumably due to a lack of vitellogenin transport out of the intestines to maturing oocytes. We used RNAi to knockdown vit-1, vit-3, vit-5, or all three genes simultaneously (vit-1,3,5) and monitored intestinal activation of the UPRmt in young adult nematodes. We used the UPRmt reporter in which the GFP expression is driven by the promoter of the mitochondrial chaperone mtHsp70/hsp-6 (Yoneda et al. 2004). We additionally used RNAi of OSCP/atp-3 to induce the UPRmt in young adult nematodes as previously described (Angeli et al. 2021). We found that in hermaphroditic nematodes, knockdown of vit-1 trended toward elevating the intestinal UPRmt (p=0.0504) and knockdown of vit-3, vit-5, or vit-1,3,5 all significantly elevated the UPRmt (Figure 1A-1D). In contrast, knockdown of vit-1, vit-3, or vit-5 did not significantly alter the UPRmt of fem-1 mutants (Figure 1A-1D). The knockdown of vit-1,3,5 did significantly enhance the UPRmt of the fem-1 mutants, but not to wild-types levels (Figure 1D). One caveat of this study is that we were not able to procure RNAi clones for vit-2, vit-4, or vit-6. However, since vit-2 is over 80% identical to vit-1 and vit-3/vit-4 are duplicated genes that are 99% identical (Perez and Lehner 2019), we would expect to observe more impact from individual RNAi knockdown on females if these vitellogenins played an outsized role on the UPRmt. The vit-6 gene is the most divergent vitellogenin, so it remains possible that it plays an untested role in the UPRmt. Overall, while it does appear that vitellogenins can mildly suppress the adult UPRmt, especially in hermaphroditic nematodes, we conclude that additional germline-to-intestinal signals are responsible for the potent suppression of the UPRmt in female nematodes.
","references":[{"reference":"Angeli S, Foulger A, Chamoli M, Peiris TH, Gerencser A, Shahmirzadi AA, Andersen J, Lithgow G. 2021. The mitochondrial permeability transition pore activates the mitochondrial unfolded protein response and promotes aging. eLife 10: 10.7554/elife.63453.
","pubmedId":"","doi":"10.7554/eLife.63453"},{"reference":"Charmpilas N, Sotiriou A, Axarlis K, Tavernarakis N, Hoppe T. 2024. Reproductive regulation of the mitochondrial stress response in Caenorhabditis elegans. Cell Reports 43: 114336.
","pubmedId":"","doi":"10.1016/j.celrep.2024.114336"},{"reference":"DePina AS, Iser WB, Park SS, Maudsley S, Wilson MA, Wolkow CA. 2011. Regulation of Caenorhabditis elegans vitellogenesis by DAF-2/IIS through separable transcriptional and posttranscriptional mechanisms. BMC Physiology 11: 10.1186/1472-6793-11-11.
","pubmedId":"","doi":"10.1186/1472-6793-11-11"},{"reference":"Dimov I, Maduro MF. 2019. The C. elegans intestine: organogenesis, digestion, and physiology. Cell and Tissue Research 377: 383-396.
","pubmedId":"","doi":" 10.1007/s00441-019-03036-4"},{"reference":"Foulger AC, Jordan NA, Castle A, Bhaumik D, Andersen JK, Lithgow GJ, Angeli S. 2025. Germline regulation of the intestinal mitochondrial unfolded protein response. GeroScience : 10.1007/s11357-025-01890-5.
","pubmedId":"","doi":"10.1007/s11357-025-01890-5"},{"reference":"Kim S, Ramalho TR, Haynes CM. 2024. Regulation of proteostasis and innate immunity via mitochondria-nuclear communication. Journal of Cell Biology 223: 10.1083/jcb.202310005.
","pubmedId":"","doi":"10.1083/jcb.202310005"},{"reference":"Perez MF, Lehner B. 2019. Vitellogenins - Yolk Gene Function and Regulation in Caenorhabditis elegans. Frontiers in Physiology 10: 10.3389/fphys.2019.01067.
","pubmedId":"","doi":"10.3389/fphys.2019.01067"},{"reference":"Shen K, Durieux J, Mena CG, Webster BM, Tsui CK, Zhang H, et al., Dillin. 2024. The germline coordinates mitokine signaling. Cell 187: 4605-4620.e17.
","pubmedId":"","doi":" 10.1016/j.cell.2024.06.010"},{"reference":"Sulston J, H.J., Methods, in The Nematode Caenorhabditis elegans, W.B. W., Editor. 1988, Cold Spring Harbor Laboratory Press Cold Spring Harbor. p. 587-606
","pubmedId":"","doi":""},{"reference":"Yoneda T, Benedetti C, Urano F, Clark SG, Harding HP, Ron D. 2004. Compartment-specific perturbation of protein handling activates genes encoding mitochondrial chaperones. Journal of Cell Science 117: 4055-4066.
","pubmedId":"","doi":" 10.1242/jcs.01275"},{"reference":"Zhou L, Jiang L, Li L, Ma C, Xia P, Ding W, Liu Y. 2024. A germline-to-soma signal triggers an age-related decline of mitochondrial stress response. Nature Communications 15: 10.1038/s41467-024-53064-0.
","pubmedId":"","doi":" 10.1038/s41467-024-53064-0"}],"title":"Role of vitellogenins on the mitochondrial unfolded protein response
","reviews":[],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":"1790462974678"}]},{"id":"ecf00b95-8783-4fa6-83d2-f24677e96cac","decision":"publish","abstract":"Sex-specific differences influence aging and disease onset. Using Caenorhabditis elegans, we previously found that hermaphroditic nematodes activate a robust mitochondrial unfolded protein response (UPRmt) in the intestines, while female nematodes do not. We hypothesized that the accumulation of vitellogenins in females was suppressing the UPRmt. Loss of vitellogenins via RNAi enhanced the UPRmt in hermaphrodites but had no impact on the UPRmt of females unless a combination of vitellogenins was simultaneously knocked down. Furthermore, combinatorial loss of vitellogenins did not restore the female UPRmt to hermaphroditic levels. We conclude that additional factors besides vitellogenins are involved in suppressing the intestinal UPRmt in female nematodes.
","acknowledgements":"We thank the Caenorhabditis Genetics Center (CGC), which is funded by the NIH Office of Research Infrastructure Programs (P40 OD010440).
","authors":[{"affiliations":["University of Maine"],"departments":["Molecular and Biomedical Sciences"],"credit":["dataCuration","methodology","validation","writing_originalDraft","formalAnalysis"],"email":"nathaniel.a.jordan@maine.edu","firstName":"Nathanial A.","lastName":"Jordan","submittingAuthor":false,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":null},{"affiliations":["University of Maine"],"departments":["Molecular and Biomedical Sciences"],"credit":["conceptualization","fundingAcquisition","supervision","writing_reviewEditing"],"email":"suzanne.angeli@maine.edu","firstName":"Suzanne","lastName":"Angeli","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[{"awardId":"5P20GM144265 - 03 ","funderName":"National Institute of General Medical Sciences (United States)","awardRecipient":"Suzanne Angeli"}],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"NIGMS COBRE Research Pilot Project Subaward 5P20GM144265 - 03 and the UMaine Institute of Medicine.
","image":{"url":"https://portal.micropublication.org/uploads/d6b2a6ea0e651800efdd64682967f9f4.png"},"imageCaption":"A-D. Quantification of GFP intensity from phsp-6::GFP or fem-1(hc17); phsp-6::GFP nematodes. For single vitellogenin gene knockdown, nematodes were developed on either control vector (CV) RNAi or vit-1 (A), vit-3 (B), or vit-5 (C) RNAi at 25°C. At the young adult stage, control worms were shifted to 2o°C and onto RNAi plates seeded with CV or CV/atp-3 (50%/50% RNAi mixture). Vitellogenin RNAi treated worms were shifted to 2o°C and onto RNAi plates seeded with the indicated CV/vit RNAi or vit/atp-3 (50%/50% RNAi mixture). For the combinatorial vitellogenin gene knockdown (D), nematodes were developed on either control vector (CV) RNAi or vit-1, vit-3, vit-5 (33%/33%/33% mixture) RNAi at 25°C. At the young adult stage, vitellogenin RNAi treated worms were shifted to 2o°C and onto RNAi plates seeded with the CV/vit-1,3,5 RNAi (50%/50% combinatorial RNAi mixture) or vit-1,3,5/atp-3 (50% combinatorial mixture/50% atp-3 RNAi). Error bars represent ± SEM. ***p ≤ 0.001, **p≤ 0.01, and *p≤ 0.05.
","imageTitle":"Impact of vitellogenin RNAi on the UPRmt
","methods":"Strains. Bristol N2 (wild type) nematodes were obtained from the Caenorhabditis Genetics Center (CGC, University of Minnesota) and cultured using standard conditions (Sulston J 1988). The following strains were used: GL347 (SJ4100 phsp-6::GFP backcrossed 6× to N2) and GL364 (fem-1(hc17); phsp-6::GFP).
Nematode and bacterial culture conditions. Nematodes were maintained on nematode growth medium (NGM) plates. NGM plates were seeded with Escherichia coli OP50 obtained from CGC that was grown in LB at 37°C for 18 hours shaking at 225 rpm. Seeded plates were dried for 48 hours at room temperature before use. For RNAi experiments, HT115 (DE3) bacteria obtained from the Horizon Biosciences RNAi library were used. All RNAi clones were verified via sequencing. RNAi plates were prepared by cooling NGM to 55°C and supplementing with a final concentration of 50μg/ml carbenicillin and 1 mM isopropyl β-d-1-thiogalactopyranoside (IPTG). RNAi bacteria were inoculated with one colony of RNAi bacteria into LB with 50μg/ml carbenicillin and were grown shaking overnight for 18 hours at 37° at 225 rpm. RNAi cultures were seeded on RNAi plates and allowed to grow for 48 hours at room temperature. Plates were stored at 4 °C for no longer than 2 weeks.
Microscopy. Nematodes were anesthetized with 2 - 5mM levamisole and mounted on 2% agarose pads on glass slides. Fluorescence micrographs of GFP were taken using a Zeiss Axioscope 5 fluorescent compound microscope equipped with Zen microscopy imaging program. GFP expression was enhanced using the brightness/contrast tool in Adobe Photoshop. The same parameters were used for all images. GFP intensity of nematodes was quantified using ImageJ 1.54G. The “integrated density” of GFP expression and length of nematodes was measured using ImageJ tools. Integrated density value was normalized by number of nematodes and average length of nematodes. The final value is in arbitrary units.
Statistics. Significance between control and experimental groups was determined by using a two-tailed Student's t-test. Asterisks denote corresponding statistical significance: *p < 0.05; **p < 0.01; ***p < 0.001. Error bars were generated using the standard error of the mean (SEM), typically from three or more pooled biological replicates.
","reagents":"","patternDescription":"Age-related diseases, such as cardiovascular disease, cancer, and neurodegenerative syndromes can display sex-specific differences. Since mitochondrial dysfunction is central to most age-related diseases, we examined the mitochondrial unfolded protein response (UPRmt), a conserved, broad-range transcriptional response that, among other functions, aids in the refolding of mitochondrial matrix proteins (Kim, Ramalho, and Haynes 2024). In C. elegans, UPRmt activation is largely localized to the intestine, the major metabolic tissue in the nematode. In addition to digestion, the intestine also carries out many functions similar to the liver, such as detoxification, immunity, and fat metabolism (Dimov and Maduro 2019). We and others recently discovered that the activation of the intestinal UPRmt depends on the function of another tissue: the germline (Foulger et al. 2025; Charmpilas et al. 2024; Shen et al. 2024; Zhou et al. 2024). Specifically, we found that adult nematodes with actively proliferating germlines, such as hermaphrodites or mated females, can activate a robust UPRmt in the intestines when challenged with either a high dose of the metal manganese (Mn) or RNAi of the OSCP/atp-3 subunit of complex V (Foulger et al. 2025). Conversely, adult nematodes with mutations that lead to a lack germline stem cells or lack of sperm, such glp-1 or fem-1 mutants, cannot activate the intestinal UPRmt. Additionally, we found that loss of the transcription factor FOXO/daf-16 fully restored the UPRmt in fem-1 mutants (Foulger et al. 2025). Here, we probe whether vitellogenesis, which occurs in the intestines and is dependent on FOXO/daf-16, impacts the UPRmt.
Vitellogenins are precursor yolk proteins that provide nutrients to developing embryos. In C. elegans, the highly conserved genes vit-1 through vit-6 are responsible for encoding these proteins (Perez and Lehner 2019). In hermaphrodites, they are synthesized in the intestines and transported into the pseudocoelom and to maturing oocytes (Perez and Lehner 2019). Young adult fem-1 mutants display elevated yolk content compared to wild-type hermaphrodites at Day 1 of adulthood (DePina et al. 2011), presumably due to a lack of vitellogenin transport out of the intestines to maturing oocytes. We used RNAi to knockdown vit-1, vit-3, vit-5, or all three genes simultaneously (vit-1,3,5) and monitored intestinal activation of the UPRmt in young adult nematodes. We used the UPRmt reporter in which the GFP expression is driven by the promoter of the mitochondrial chaperone mtHsp70/hsp-6 (Yoneda et al. 2004). We additionally used RNAi of OSCP/atp-3 to induce the UPRmt in young adult nematodes as previously described (Angeli et al. 2021). We found that in hermaphroditic nematodes, knockdown of vit-1 trended toward elevating the intestinal UPRmt (p=0.0504) and knockdown of vit-3, vit-5, or vit-1,3,5 all significantly elevated the UPRmt (Figure 1A-1D). In contrast, knockdown of vit-1, vit-3, or vit-5 did not significantly alter the UPRmt of fem-1 mutants (Figure 1A-1D). The knockdown of vit-1,3,5 did significantly enhance the UPRmt of the fem-1 mutants, but not to wild-types levels (Figure 1D). One caveat of this study is that we were not able to procure RNAi clones for vit-2, vit-4, or vit-6. However, since vit-2 is over 80% identical to vit-1 and vit-3/vit-4 are duplicated genes that are 99% identical (Perez and Lehner 2019), we would expect to observe more impact from individual RNAi knockdown on females if these vitellogenins played an outsized role on the UPRmt. The vit-6 gene is the most divergent vitellogenin, so it remains possible that it plays an untested role in the UPRmt. Overall, while it does appear that vitellogenins can mildly suppress the adult UPRmt, especially in hermaphroditic nematodes, we conclude that additional germline-to-intestinal signals are responsible for the potent suppression of the UPRmt in female nematodes.
","references":[{"reference":"Angeli S, Foulger A, Chamoli M, Peiris TH, Gerencser A, Shahmirzadi AA, Andersen J, Lithgow G. 2021. The mitochondrial permeability transition pore activates the mitochondrial unfolded protein response and promotes aging. eLife 10: 10.7554/elife.63453.
","pubmedId":"","doi":"10.7554/eLife.63453"},{"reference":"Charmpilas N, Sotiriou A, Axarlis K, Tavernarakis N, Hoppe T. 2024. Reproductive regulation of the mitochondrial stress response in Caenorhabditis elegans. Cell Reports 43: 114336.
","pubmedId":"","doi":"10.1016/j.celrep.2024.114336"},{"reference":"DePina AS, Iser WB, Park SS, Maudsley S, Wilson MA, Wolkow CA. 2011. Regulation of Caenorhabditis elegans vitellogenesis by DAF-2/IIS through separable transcriptional and posttranscriptional mechanisms. BMC Physiology 11: 10.1186/1472-6793-11-11.
","pubmedId":"","doi":"10.1186/1472-6793-11-11"},{"reference":"Dimov I, Maduro MF. 2019. The C. elegans intestine: organogenesis, digestion, and physiology. Cell and Tissue Research 377: 383-396.
","pubmedId":"","doi":" 10.1007/s00441-019-03036-4"},{"reference":"Foulger AC, Jordan NA, Castle A, Bhaumik D, Andersen JK, Lithgow GJ, Angeli S. 2025. Germline regulation of the intestinal mitochondrial unfolded protein response. GeroScience : 10.1007/s11357-025-01890-5.
","pubmedId":"","doi":"10.1007/s11357-025-01890-5"},{"reference":"Kim S, Ramalho TR, Haynes CM. 2024. Regulation of proteostasis and innate immunity via mitochondria-nuclear communication. Journal of Cell Biology 223: 10.1083/jcb.202310005.
","pubmedId":"","doi":"10.1083/jcb.202310005"},{"reference":"Perez MF, Lehner B. 2019. Vitellogenins - Yolk Gene Function and Regulation in Caenorhabditis elegans. Frontiers in Physiology 10: 10.3389/fphys.2019.01067.
","pubmedId":"","doi":"10.3389/fphys.2019.01067"},{"reference":"Shen K, Durieux J, Mena CG, Webster BM, Tsui CK, Zhang H, et al., Dillin. 2024. The germline coordinates mitokine signaling. Cell 187: 4605-4620.e17.
","pubmedId":"","doi":" 10.1016/j.cell.2024.06.010"},{"reference":"Sulston J, H.J., Methods, in The Nematode Caenorhabditis elegans, W.B. W., Editor. 1988, Cold Spring Harbor Laboratory Press Cold Spring Harbor. p. 587-606
","pubmedId":"","doi":""},{"reference":"Yoneda T, Benedetti C, Urano F, Clark SG, Harding HP, Ron D. 2004. Compartment-specific perturbation of protein handling activates genes encoding mitochondrial chaperones. Journal of Cell Science 117: 4055-4066.
","pubmedId":"","doi":" 10.1242/jcs.01275"},{"reference":"Zhou L, Jiang L, Li L, Ma C, Xia P, Ding W, Liu Y. 2024. A germline-to-soma signal triggers an age-related decline of mitochondrial stress response. Nature Communications 15: 10.1038/s41467-024-53064-0.
","pubmedId":"","doi":" 10.1038/s41467-024-53064-0"}],"title":"Role of vitellogenins on the mitochondrial unfolded protein response
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