{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T18:44:27Z","timestamp":1760035467499,"version":"build-2065373602"},"reference-count":39,"publisher":"MDPI AG","issue":"7","license":[{"start":{"date-parts":[[2025,7,9]],"date-time":"2025-07-09T00:00:00Z","timestamp":1752019200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"EU Commission under the Marie Sklodowska-Curie Action\u2014European Training Network","award":["859891"],"award-info":[{"award-number":["859891"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Insects"],"abstract":"<jats:p>Springtails are adapted to life in the pore space of soil, where humidity in moist soil is close to saturation. Drought is the most important limiting factor for springtails; however, their molecular and physiological adaptations to low humidity are not well understood. The present study explored the global proteomic drought response of the springtail, Folsomia candida (Isotomidae, Collembola). In relatively dry soil (\u2212360 kPa), adult springtails initially lost body water but re-established the normal body water content over the following two weeks. Nano LC\u2013MS\/MS analysis identified a total of 1729 unique proteins. Proteomic analysis and pathway enrichment found that the proteome generally did not show a dramatic induction of proteins in response to drought stress. After an initial down-regulation of pathways related to metabolism and growth, these pathways gradually returned to the same levels as in moist soil. Other pathways such as the cytoskeleton pathway, which is important in cell proliferation and differentiation, were predominantly down-regulated throughout the experiment in drought-exposed animals, which correlated with essentially no somatic growth of the springtails in dry soil. This study facilitates the understanding of the consequences of climate change on soil functioning and fertility.<\/jats:p>","DOI":"10.3390\/insects16070707","type":"journal-article","created":{"date-parts":[[2025,7,11]],"date-time":"2025-07-11T13:44:19Z","timestamp":1752241459000},"page":"707","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Proteomic Responses of the Springtail Folsomia candida to Drought"],"prefix":"10.3390","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1887-4315","authenticated-orcid":false,"given":"Yang","family":"Wang","sequence":"first","affiliation":[{"name":"Department of Ecoscience, Aarhus University, C.F. M\u00f8llers All\u00e9 4, 8000 Aarhus, Denmark"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Stine","family":"Slotsbo","sequence":"additional","affiliation":[{"name":"Department of Ecoscience, Aarhus University, C.F. M\u00f8llers All\u00e9 4, 8000 Aarhus, Denmark"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-9873-5645","authenticated-orcid":false,"given":"Steffen Y.","family":"Bak","sequence":"additional","affiliation":[{"name":"International Flavors & Fragrances Inc., Edwin Rahrs Vej 38, 8220 Brabrand, Denmark"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0921-4796","authenticated-orcid":false,"given":"Christopher J.","family":"Martyniuk","sequence":"additional","affiliation":[{"name":"Center for Environmental and Human Toxicology, University of Florida, 2187 Mowry Road, Gainesville, FL 32611, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8395-6582","authenticated-orcid":false,"given":"Martin","family":"Holmstrup","sequence":"additional","affiliation":[{"name":"Department of Ecoscience, Aarhus University, C.F. M\u00f8llers All\u00e9 4, 8000 Aarhus, Denmark"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2025,7,9]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"674","DOI":"10.1038\/s41467-023-36216-6","article-title":"Globally invariant metabolism but density-diversity mismatch in springtails","volume":"14","author":"Potapov","year":"2023","journal-title":"Nat. Commun."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1163\/187631270X00366","article-title":"Alimentary studies of seven common soil-inhabiting Collembola of southern Sweden","volume":"1","year":"1970","journal-title":"Entomol. Scand."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1016\/j.pedobi.2011.02.004","article-title":"Tracking Collembola feeding strategies by the natural C-13 signal of fatty acids in an arable soil with different fertilizer regimes","volume":"54","author":"Ngosong","year":"2011","journal-title":"Pedobiologia"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1016\/j.soilbio.2016.07.002","article-title":"Connecting taxonomy and ecology: Trophic niches of collembolans as related to taxonomic identity and life forms","volume":"101","author":"Potapov","year":"2016","journal-title":"Soil Biol. Biochem."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1007","DOI":"10.1016\/S0038-0717(99)00014-0","article-title":"Experimental evidence that soil fauna enhance nutrient mineralization and plant nutrient uptake in montane grassland ecosystems","volume":"31","author":"Bardgett","year":"1999","journal-title":"Soil Biol. Biochem."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"565","DOI":"10.5194\/soil-2-565-2016","article-title":"Soil fauna: Key to new carbon models","volume":"2","author":"Filser","year":"2016","journal-title":"Soil"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"2727","DOI":"10.1111\/gcb.14685","article-title":"Fast attrition of springtail communities by experimental drought and richness-decomposition relationships across Europe","volume":"25","author":"Peguero","year":"2019","journal-title":"Glob. Change Biol."},{"key":"ref_8","first-page":"39","article-title":"The porosphere as an ecological medium emphasized in Professor Ghilarov\u2019s work on soil animal adaptations","volume":"3","author":"Vannier","year":"1987","journal-title":"Biol. Fertil. Soils"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1007\/s00442-011-1909-0","article-title":"A meta-analysis of responses of soil biota to global change","volume":"165","author":"Blankinship","year":"2011","journal-title":"Oecologia"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1961","DOI":"10.1111\/ele.14068","article-title":"Responses of soil fauna communities to the individual and combined effects of multiple global change factors","volume":"25","author":"Peng","year":"2022","journal-title":"Ecol. Lett."},{"key":"ref_11","unstructured":"EEA (2017). Climate Change, Impacts and Vulnerability in Europe 2016, European Environment Agency. An Indicator-Based Report."},{"key":"ref_12","unstructured":"P\u00f6rtner, H.-O., Roberts, D.C., Tignor, M., Poloczanska, E.S., Mintenbeck, K., Alegr\u00eda, A., Craig, M., Langsdorf, S., L\u00f6schke, S., and M\u00f6ller, V. (2022). Impacts, Adaptation, and Vulnerability. Contribution of Working Group II to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Climate Change 2022, Cambridge University Press."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1550","DOI":"10.1111\/1365-2435.13745","article-title":"Water availability rather than temperature control soil fauna community structure and prey-predator interactions","volume":"35","author":"Baldy","year":"2021","journal-title":"Funct. Ecol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"229","DOI":"10.14411\/eje.2019.025","article-title":"Water relations and drought sensitivity of Folsomia candida eggs (Collembola: Isotomidae)","volume":"116","author":"Holmstrup","year":"2019","journal-title":"Eur. J. Entomol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"115913","DOI":"10.1016\/j.geoderma.2022.115913","article-title":"Soil dwelling springtails are resilient to extreme drought in soil, but their reproduction is highly sensitive to small decreases in soil water potential","volume":"421","author":"Wang","year":"2022","journal-title":"Geoderma"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1909","DOI":"10.1126\/science.285.5435.1909","article-title":"Water vapor absorption in arthropods by accumulation of myoinositol and glucose","volume":"285","author":"Bayley","year":"1999","journal-title":"Science"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1046\/j.0269-8463.2001.00565.x","article-title":"Dehydration tolerance and water vapour absorption in two species of soil-dwelling Collembola by accumulation of sugars and polyols","volume":"15","author":"Holmstrup","year":"2001","journal-title":"Funct. Ecol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1021","DOI":"10.1016\/S0022-1910(01)00078-6","article-title":"Enhanced drought tolerance of a soil-dwelling springtail by pre-acclimation to a mild drought stress","volume":"47","author":"Sjursen","year":"2001","journal-title":"J. Insect Physiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"939","DOI":"10.1007\/s00360-018-1180-3","article-title":"The springtail Megaphorura arctica survives extremely high osmolality of body fluids during drought","volume":"188","author":"Holmstrup","year":"2018","journal-title":"J. Comp. Physiol. B"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1104","DOI":"10.1016\/j.jinsphys.2013.08.015","article-title":"Protaphorura tricampata, a euedaphic and highly permeable springtail that can sustain activity by osmoregulation during extreme drought","volume":"59","author":"Holmstrup","year":"2013","journal-title":"J. Insect Physiol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1016\/j.jinsphys.2015.09.005","article-title":"Accumulation of free amino acids during exposure to drought in three springtail species","volume":"82","author":"Holmstrup","year":"2015","journal-title":"J. Insect Physiol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Clark, M.S., Thorne, M.A.S., Purac, J., Burns, G., Hillyard, G., Popovic, Z.D., Grubor-Lajsic, G., and Worland, M.R. (2009). Surviving the cold: Molecular analyses of insect cryoprotective dehydration in the Arctic springtail Megaphorura arctica (Tullberg). BMC Genom., 10.","DOI":"10.1186\/1471-2164-10-328"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"737","DOI":"10.1111\/j.1365-2583.2009.00916.x","article-title":"Sugar sweet springtails: On the transcriptional response of Folsomia candida (Collembola) to desiccation stress","volume":"18","author":"Timmermans","year":"2009","journal-title":"Insect Mol. Biol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1111\/j.1365-2583.2010.01062.x","article-title":"Proteomics of cryoprotective dehydration in Megaphorura arctica Tullberg 1876 (Onychiuridae: Collembola)","volume":"20","author":"Thorne","year":"2011","journal-title":"Insect Mol. Biol."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Faddeeva-Vakhrusheva, A., Kraaijeveld, K., Derks, M.F.L., Anvar, S.Y., Agamennone, V., Suring, W., Kampfraath, A.A., Ellers, J., Le Ngoc, G., and van Gestel, C.A.M. (2017). Coping with living in the soil: The genome of the parthenogenetic springtail Folsomia candida. BMC Genom., 18.","DOI":"10.1186\/s12864-017-3852-x"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1111\/j.1365-2435.2009.01541.x","article-title":"Dynamics of heat-induced thermal stress resistance and Hsp70 expression in the springtail, Orchesella cincta","volume":"23","author":"Bahrndorff","year":"2009","journal-title":"Funct. Ecol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1006","DOI":"10.1002\/pmic.201300553","article-title":"Suspension trapping (STrap) sample preparation method for bottom- up proteomics analysis","volume":"14","author":"Zougman","year":"2014","journal-title":"Proteomics"},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Kumar, R., Kane, H., Wang, Q., Hibberd, A., Jensen, H.M., Kim, H.S., Bak, S.Y., Auzanneau, I., Bry, S., and Christensen, N. (2022). Identification and characterization of a novel species of genus Akkermansia with metabolic health effects in a diet-induced obesity mouse model. Cells, 11.","DOI":"10.3390\/cells11132084"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"923","DOI":"10.1038\/nmeth1113","article-title":"Semi-supervised learning for peptide identification from shotgun proteomics datasets","volume":"4","author":"Kall","year":"2007","journal-title":"Nat. Methods"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1074\/mcp.M500230-MCP200","article-title":"Absolute quantification of proteins by LCMSE: A virtue of parallel MS acquisition","volume":"5","author":"Silva","year":"2006","journal-title":"Mol. Cell Proteom."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1093\/bioinformatics\/btp616","article-title":"edgeR: A Bioconductor package for differential expression analysis of digital gene expression data","volume":"26","author":"Robinson","year":"2010","journal-title":"Bioinformatics"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"2565","DOI":"10.1002\/pmic.201100033","article-title":"The role of statistical power analysis in quantitative proteomics","volume":"11","author":"Levin","year":"2011","journal-title":"Proteomics"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1089\/omi.2011.0118","article-title":"clusterProfiler: An R package for comparing biological themes among gene clusters","volume":"16","author":"Yu","year":"2012","journal-title":"Omics-A J. Integr. Biol."},{"key":"ref_34","first-page":"159","article-title":"Metabolic responses to anabiosis in the 4th stage juveniles of Ditylenchus dipsaci (nematoda)","volume":"216","author":"Barrett","year":"1982","journal-title":"Proc. R. Soc. Ser. B-Biol. Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1086\/651459","article-title":"Metabolic changes during estivation in the common earthworm Aporrectodea caliginosa","volume":"83","author":"Bayley","year":"2010","journal-title":"Physiol. Biochem. Zool."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1002\/jez.1402010107","article-title":"Anhydrobiosis in nematodes: Metabolism during resumption of activity","volume":"201","author":"Crowe","year":"1977","journal-title":"J. Exp. Zool."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Gleason, L.U., Fekete, F.J., Tanner, R.L., and Dowd, W.W. (2023). Multi-omics reveals largely distinct transcript- and protein-level responses to the environment in an intertidal mussel. J. Exp. Biol., 226.","DOI":"10.1242\/jeb.245962"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1146\/annurev.arplant.54.031902.134818","article-title":"Remodeling the cytoskeleton for growth and form: An overview with some new views","volume":"54","author":"Wasteneys","year":"2003","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"961","DOI":"10.1016\/S0022-1910(02)00175-0","article-title":"Drought acclimation and lipid composition in Folsomia candida: Implications for cold shock, heat shock and acute desiccation stress","volume":"48","author":"Holmstrup","year":"2002","journal-title":"J. Insect Physiol."}],"container-title":["Insects"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2075-4450\/16\/7\/707\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T18:07:29Z","timestamp":1760033249000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2075-4450\/16\/7\/707"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,9]]},"references-count":39,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2025,7]]}},"alternative-id":["insects16070707"],"URL":"https:\/\/doi.org\/10.3390\/insects16070707","relation":{},"ISSN":["2075-4450"],"issn-type":[{"type":"electronic","value":"2075-4450"}],"subject":[],"published":{"date-parts":[[2025,7,9]]}}}