Publications

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Patrício Silva, A. L., Holmstrup, M. & Amorim, M. J. B. (2013). Worms from the Arctic are better adapted to freezing and high salinity than worms from temperate regions: Oxidative stress responses in Enchytraeus albidus. Comparative Biochemistry and Physiology -Part A : Molecular and Integrative Physiology, 166(4), 582-589. https://doi.org/10.1016/j.cbpa.2013.09.004

Slotsbo, S., Hansen, L. M. & Holmstrup, M. (2010). Winter survival of the Iberian slug Arion lusitanicus. In J. Kollmann, T. van Mölken & H. P. Ravn (Eds.), Biological Invasions in a Changing World from Science to Management. : Book of Abstracts (pp. 10)

Krogh, P. H., Jensen, J., Holmstrup, M., Scott-Fordsmand, J. J. & Kristensen, H. L. (1999). Why is Sludge with Toxicant Load Non-Toxic in the Field?. Poster session presented at Annual Meeting of SETAC-Europe, Germany.

Naveed, M., Møldrup, P., Arthur, E., Holmstrup, M., Nicolaisen, M., Herath, L., Hamamoto, S., Kawamoto, K., Komatsu, T., Tuller, M. & de Jonge, L. W. (2013). When Soil Goes to Sleep: Simultaneous Changes in Biodiversity and Soil Ecosystem Functions Along an Anthropogenic Copper Gradient. In Water, Food, Energy & Innovation for a Sustainable World Article 289-9 https://scisoc.confex.com/crops/2013am/webprogram/Paper81623.html

Bayley, M. & Holmstrup, M. (1999). Water Vapor Absorption in Arthropods by Accumulation of Myoinositol and Glucose. Science, 285(5435), 1909-1911. https://doi.org/10.1126/science.285.5435.1909

Holmstrup, M. (2019). Water relations and drought sensitivity of Folsomia candida eggs (Collembola: Isotomidae). European Journal of Entomology, 116, 229-234. Article 025. https://doi.org/10.14411/eje.2019.025

Thakur, M. P., Sigurðsson, B. D., Sigurðsson, P. & Holmstrup, M. (2023). Warming shifts the biomass distribution of soil microarthropod communities. Soil Biology and Biochemistry, 177, Article 108894. https://doi.org/10.1016/j.soilbio.2022.108894

Fisker, K. V., Holmstrup, M. & Sørensen, J. G. (2013). Variation in metallothionein gene expression is associated with adaptation to copper in the earthworm Dendrobaena octaedra. Comparative Biochemistry and Physiology Part - C: Toxicology and Pharmacology, 157(2), 220-226. https://doi.org/10.1016/j.cbpc.2012.11.007

Rasmussen, L. B., Jensen, K., Sørensen, J. G., Toft, S., Overgaard, J., Holmstrup, M. & Kristensen, T. N. (2017). Variable performance and improvement by crossing in commercial populations of the pirate bug Orius majusculus. Poster session presented at 5th International Symposium on Biological Control of Arthropods, Langkawi, Malaysia.

Rasmussen, L. B., Jensen, K., Sørensen, J. G., Overgaard, J., Holmstrup, M. & Kristensen, T. N. (2017). Variable performance and improvement by crossing in commercial populations of the pirate bug Orius majusculus. In P. G. Mason, D. R. Gillespie & C. Vincent (Eds.), Proceedings of the 5th international symposium on biological control of arthropods (pp. 243). CABI Publishing.

Rasmussen, L. B., Jensen, K., Sørensen, J. G., Toft, S., Overgaard, J., Holmstrup, M. & Kristensen, T. N. (2017). Variable performance and improvement by crossing in commercial populations of the pirate bug Orius majusculus. Abstract from 5th International Symposium on Biological Control of Arthropods, Langkawi, Malaysia.

Schmidt, S. N., Smith, K. E. C., Holmstrup, M. & Mayer, P. (2013). Uptake and toxicity of polycyclic aromatic hydrocarbons in terrestrial springtails—studying bioconcentration kinetics and linking toxicity to chemical activity. Environmental Toxicology and Chemistry, 32(2), 361-369. https://doi.org/10.1002/etc.2051

Patrício Silva, A. L., Amorim, M. J. B. & Holmstrup, M. (2016). Uptake and Elimination of 4-Nonylphenol in the EnchytraeidEnchytraeus albidus. Bulletin of Environmental Contamination and Toxicology, 96(2), 156–161. https://doi.org/10.1007/s00128-015-1701-7

Slotsbo, S., Sørensen, J. G., Holmstrup, M., Kostál, V., Kellermann, V. & Overgaard, J. (2016). Tropical to subpolar gradient in phospholipid composition suggests adaptive tuning of biological membrane function in drosophilids. Functional Ecology, 30(5), 759-768. https://doi.org/10.1111/1365-2435.12568

Peng, Y., Holmstrup, M., Kappel Schmidt, I., Ruggiero Bachega, L., Schelfhout, S., Zheng, H., Heděnec, P., Yue, K. & Vesterdal, L. (2022). Tree species identity is the predominant modulator of the effects of soil fauna on leaf litter decomposition. Forest Ecology and Management, 520, Article 120396. https://doi.org/10.1016/j.foreco.2022.120396

Paul, S., Heckmann, L.-H., Sorensen, J. G., Holmstrup, M., Arumugaperumal, A. & Sivasubramaniam, S. (2018). Transcriptome sequencing, de novo assembly and annotation of the freeze tolerant earthworm, Dendrobaena octaedra. Gene Reports, 13, 180-191. https://doi.org/10.1016/j.genrep.2018.10.010

Orr, J. A., Vinebrooke, R. D., Jackson, M. C., Kroeker, K. J., Kordas, R. L., Mantyka-Pringle, C., Van den Brink, P. J., De Laender, F., Stoks, R., Holmstrup, M., Matthaei, C. D., Monk, W. A., Penk, M. R., Leuzinger, S., Schäfer, R. B. & Piggott, J. J. (2020). Towards a unified study of multiple stressors: divisions and common goals across research disciplines. Proceedings of the Royal Society B: Biological Sciences, 287(1926), Article 20200421. https://doi.org/10.1098/rspb.2020.0421

Løkke, H., Ragas, A. M. J. & Holmstrup, M. (2013). Tools and perspectives for assessing chemical mixtures and multiple stressors. Toxicology, 313, 73-82. https://doi.org/10.1016/j.tox.2012.11.009

Xie, L., Slotsbo, S. & Holmstrup, M. (2023). Tolerance of high temperature and associated effects on reproduction in euedaphic Collembola. Journal of Thermal Biology, 113, Article 103439. https://doi.org/10.1016/j.jtherbio.2022.103439

Grantland, D., Holmstrup, M., Overgaard, J., Holmstrup, M. E. & Slotsbo, S. (2025). Tiny Worms, Big Potential: Enchytraeus albidus (Annelida: Clitellata) as Starter Feed for Rainbow Trout. Aquaculture Research, 2025(1), Article 7339644. https://doi.org/10.1155/are/7339644

Coulson, S., Convey, P., Aakra, K., Aarvik, L., Ávila-Jiménez, M. L., Babenko, A., Biersma, E. M., Boström, S., Brittain, J. E., Carlsson, A. M., Christoffersen, K., De Smet, W. H., Ekremj, T., Fjellberg, A., Füreder, L., Gustafsson, D., Gwiazdowicz, D. J., Hansen, L. O., Holmstrup, M. ... Zmudczynska-Skarbek, K. (2014). The terrestrial and freshwater invertebrate biodiversity of the archipelagoes of the Barents Sea; Svalbard, Franz Josef Land and Novaya Zemlya. Soil Biology & Biochemistry, 68, 440-470. https://doi.org/10.1016/j.soilbio.2013.10.006

Holmstrup, M. (2018). The springtail Megaphorura arctica survives extremely high osmolality of body fluids during drought. Journal of Comparative Physiology B: Biochemical, Systems, and Environmental Physiology, 188(6), 939-945. https://doi.org/10.1007/s00360-018-1180-3

Nasiri Moghadam, N., Holmstrup, M., Manenti, T., Mouridsen, M., Pertoldi, C. & Loeschcke, V. (2015). The Role of Storage Lipids in the Relation between Fecundity, Locomotor Activity, and Lifespan of Drosophila melanogaster Longevity-Selected and Control Lines. PLoS One, 10(6), Article e0130334. https://doi.org/10.1371/journal.pone.0130334

Engell Dahl, J., Marti, S. L., Colinet, H., Wiegand, C., Holmstrup, M. & Renault, D. (2021). Thermal plasticity and sensitivity to insecticides in populations of an invasive beetle: Cyfluthrin increases vulnerability to extreme temperature. Chemosphere, 274, Article 129905. https://doi.org/10.1016/j.chemosphere.2021.129905

Dai, W., Slotsbo, S. & Holmstrup, M. (2021). Thermal optimum for mass production of the live feed organism Enchytraeus albidus. Journal of Thermal Biology, 97, Article 102865. https://doi.org/10.1016/j.jtherbio.2021.102865

Bahrndorff, S., Alemu, T., Kristensen, T. N., Sørensen, M. H., Høye, T. T. & Holmstrup, M. (2021). Thermal adaptations of adults and eggs in the Arctic seed bug Nysius groenlandicus (Insecta: Hemiptera) from South Greenland. Polar Biology, 44(3), 491-498. https://doi.org/10.1007/s00300-021-02807-6

Bahrndorff, S., Loeschcke, V., Pertoldi, C., Beier, C. & Holmstrup, M. (2009). The rapid cold hardening response of Collembola is influenced by thermal variability of the habitat. Functional Ecology, 23(2), 340–347. https://doi.org/10.1111/j.1365-2435.2008.01503.x

Holmstrup, M. (2014). The ins and outs of water dynamics in cold tolerant soil invertebrates. Journal of Thermal Biology, 45, 117-123. https://doi.org/10.1016/j.jtherbio.2014.09.001

Slotsbo, S., Hansen, L. M., Damgaard, C. & Holmstrup, M. (2013). The influence of temperature on life history traits in the Iberian slug, Arion lusitanicus. Annals of Applied Biology, 162, 80-88.

Widarto, T. H., Holmstrup, M. & Forbes, V. E. (2004). The influence of nonylphenol on the life-history of the earthworm Dendrobaena octaedra Savigny: Linking effects from the individual to the population level. Ecotoxicology and Environmental Safety, 58, 147-159.

Kærsgaard, C. W., Holmstrup, M., Malte, H. & Bayley, M. (2004). The importance of cuticular permeability, osmolyte production and body size for the desiccation resistance of nine species of Collembola. Journal of Insect Physiology, 50(1), 5-15. https://doi.org/10.1016/j.jinsphys.2003.09.003

Maraldo, K., Christensen, B. & Holmstrup, M. (2011). The excretion of ammonium by enchytraeids (Cognettia sphagnetorum). Soil Biology & Biochemistry, 43(5), 991-996. https://doi.org/10.1016/j.soilbio.2011.01.015

de Boer, T. E., Holmstrup, M., van Straalen, N. M. & Roelofs, D. (2010). The effect of soil pH and temperature on Folsomia candida transcriptional regulation. Journal of Insect Physiology, 56(4), 350-355. https://doi.org/10.1016/j.jinsphys.2009.11.004

Damgaard, C., Strandberg, B., Dupont, Y., Holmstrup, M. & Krogh, P. H. (2016). The effect of glyphosate and nitrogen on plant communities and the soil fauna in terrestrial biotopes at field margins. The Danish Environmental Protection Agency. Bekæmpelsesmiddelforskning fra Miljøstyrelsen No. 163 http://www2.mst.dk/Udgiv/publications/2016/09/978-87-93283-85-5.pdf

Maraldo, K., Krogh, P. H., van der Linden, L., Christensen, B., Mikkelsen, T. N., Beier, C. & Holmstrup, M. (2010). The counteracting effects of elevated atmospheric CO₂concentrations and drought episodes: Studies of enchytraeid communities in a dry heathland. Soil Biology & Biochemistry, 42(11), 1958-1966. https://doi.org/10.1016/j.soilbio.2010.07.015

Boas, S. W., Slotsbo, S. & Holmstrup, M. (2015). The combined effect of freeze thaw events and heavy metal pollution leads to distinct lethal synergy in Enchytraeus albidus. Poster session presented at ISEPEP6 - Sixth International Symposium on the Enviromental Physiology of Ectotherms and Plants, Aarhus, Denmark.

Sørensen, J. G., Heckmann, L.-H. & Holmstrup, M. (2010). Temporal gene expression profiles in a palaearctic springtail as induced by desiccation, cold exposure and during recovery. Functional Ecology, 24(4), 838-846. https://doi.org/10.1111/j.1365-2435.2010.01687.x

Kutcherov, D., Slotsbo, S., Sigurdsson, B. D., Leblans, N. I. W., Berg, M. P., Ellers, J., Mariën, J. & Holmstrup, M. (2020). Temperature responses in a subarctic springtail from two geothermally warmed habitats. Pedobiologia, 78, Article 150606. https://doi.org/10.1016/j.pedobi.2019.150606

Overgaard, J., Bayley, M. & Holmstrup, M. (2007). Temperature, humidity and inoculative freezing are all signals for glucose accumulation in the freeze tolerant earthworm Dendrobaena octaedra. Poster session presented at SEBatGlasgow 2007. Annual Main Meeting, Glasgow, United Kingdom.

Petersen, S. O. & Holmstrup, M. (2000). Temperature Effects on Lipid Composition on the Earthworms Lumbricus rubellus and Eisenia nordenskioeldi. Soil Biology & Biochemistry, 32, 1787-1791.

Marty, A., Boeriis, T., Martínez-De León, G., Holmstrup, M. & Thakur, M. P. (2022). Temperature-dependent trade-offs in maternal investments: An experimental test with two closely related soil microarthropods. European Journal of Soil Biology, 110, Article 103402. https://doi.org/10.1016/j.ejsobi.2022.103402

Dai, W., Slotsbo, S., van Gestel, C. A. M. & Holmstrup, M. (2021). Temperature-Dependent Toxicokinetics of Phenanthrene in Enchytraeus albidus (Oligochaeta). Environmental Science & Technology, 55(3), 1876-1884. https://doi.org/10.1021/acs.est.0c06182

Dai, W., Slotsbo, S., Damgaard, C., Ke, X., Wu, L. & Holmstrup, M. (2019). Synergistic interaction between effects of phenanthrene and dynamic heat stress cycles in a soil arthropod. Environmental Pollution, 254(Pt B), Article 113071. https://doi.org/10.1016/j.envpol.2019.113071

Waagner, D., Bayley, M. & Holmstrup, M. (2009). Synergistic interaction between 4-nonylphenol and high but not lowtemperatures in Dendrobaena octaedra. Ecotoxicology and Environmental Safety, 72, 10– 16.

Jensen, D., Bayley, M. & Holmstrup, M. (2009). Synergistic interaction between 4-nonylphenol and high but not low temperatures in Dendrobaena octaedra. Ecotoxicology and Environmental Safety, 72(1), 10-16. https://doi.org/10.1016/j.ecoenv.2008.03.002

Jensen, K., Toft, S., Sørensen, J. G. & Holmstrup, M. (2022). Survival and predation rate of wild-caught and commercially produced Orius majusculus (Reuter) (Hemiptera: Anthocoridae). Bulletin of Entomological Research, 112(3), 311-317. https://doi.org/10.1017/S0007485321000055

Voituron, Y., Paaschburg, L., Holmstrup, M., Barré, H. & Ramløv, H. (2009). Survival and metabolism of Rana arvalis during freezing. Journal of Comparative Physiology B: Biochemical, Systems, and Environmental Physiology, 179(2), 223-230. https://doi.org/10.1007/s00360-008-0307-3

Holmstrup, M., Bayley, M. & Ramløv, H. (2002). Supercool or Dehydrate? An Experimental Analysis of Overwintering Strategies in Small Permeable Arctic Invertebrates. Proceedings of the National Academy of Sciences (PNAS), 99(8), 5716-5720. https://doi.org/10.1073/pnas.082580699

Timmermans, M. J. T. N., Roelofs, D., Nota, B., Ylstra, B. & Holmstrup, M. (2009). Sugar sweet springtails: on the transcriptional response of Folsomia candida (Collembola) to desiccation stress. Insect Molecular Biology, 18(6), 737–746. https://doi.org/10.1111/j.1365-2583.2009.00916.x

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Revised 20.01.2026

Stine Slotsbo