Sarraude T, Hsu B-Y, Ruuskanen S, Groothuis TGG. Is maternal thyroid hormone deposition subject to a trade-off between self and egg
because of iodine? An experimental study in rock pigeon. BioRxiv, submitted to Journal of Experimental Biology

Laine VN, Verschuuren M, van Oers K, Espin S, Sanchez-Virosta P, Eeva T, Ruuskanen S*. Does arsenic contamination affect DNA methylation patterns in a wild bird population? An experimental approach. BioRxiv, submitted to Environmental Science and Technology

Mäkinen, H., van Oers, K., Eeva, T., Laine V.N., Ruuskanen, S*. The effect of experimental lead pollution on DNA methylation patterns in a wild bird population. BioRxiv, in revision for Evolutionary Applications

Hsu, B-Y., Pakanen V-M., Boner, W., Eeva, T., Doligez, B., Groothuis, TGG., Korpimäki E., Laaksonen, T., Lelono, A., Monaghan, P., Sarraude T., Tschirren, B., Thomson, RL., Tolvanen, J., Vásquez, RA., Ruuskanen, S*. Inter-specific variation of prenatal maternal thyroid hormones associate with migration, mode of development, and captivity: a phylogenetic comparative analysis. BioRxiv, in revision for Journal of Animal Ecology



[55] Rainio MJ, Ruuskanen S, Helander M, Saikkonen K, Saloniemi I, Puigbo P. 2021. Adaptation of bacteria to glyphosate: a microevolutionary perspective of the enzyme 5-enolpyruvylshikimate-3-phosphate (EPSP) synthase. Environmental Microbiology Reports, in press.

[54] Eeva T, Raivikko N, Espín S, Sanchez-Virosta P, Ruuskanen S, Sorvari J, Rainio M.2020. Bird feces as indicators of metal pollution: pitfalls and solutions. Toxics 8, 124.

[53] Leino L, Tall T, Helander M, Saloniemi I, Saikkonen K, Ruuskanen S, Puigbo P. 2021. Classification of  glyphosate’s target enzyme (5-enolpyruvylshikimate-3-phosphate synthase). Journal of Hazardous Materials (IF 9.0), in press.

Media coverage: Science Daily, Forbes, Award winner Carey Gillam:  New glyphosate papers point to “urgency” for more research on chemical impact to human health – U.S. Right to Know (

[52] Stier A, Hsu Bin-Yan, Marciau C, Doligez B, Gustafsson L, Bize P, Ruuskanen S*. Born to be young: prenatal thyroid hormones increase early-life telomere length. Biology Letters 16,20200364

Featured in: Press release, Science daily, UPI

[51] Sarraude T, Hsu BY, Groothuis TGG, Ruuskanen S*.2020. Elevated exposure to prenatal thyroid hormones affects embryonic mortality but has no effects into adulthood. PeerJ 8,e10175

[50] Sarraude T, Hsu Bin-Yan, Groothuis TGG, Ruuskanen S*. 2020.Testing different forms of regulation of yolk thyroid hormone transfer in pied flycatchers. Journal of Experimental Biology 223, jeb226688.

[49] Taborsky B, English S, Fawcett TW, Kuijper B, Leimar O, McNamara JMc, Ruuskanen S, Sandi C. 2021 Towards an Evolutionary Theory of Stress  Responses. Trends in Ecology and Evolution 36, 39-48.

Media coverage:, EureK Alert, Phys Org, UTU press release

[48] Ruuskanen S, Mottola G, Anttila K. 2020.Experimental copper exposure, but not heat stress, leads to elevated intraovarian thyroid hormone levels in three-spined sticklebacks (Gasterosteus aculeatus). Ecotoxicology, 29(9), 1431-1440.

[47] Hsu B-Y, Sarraude T, Cossin-Sevrin N, Crombecque M, Stier A* & Ruuskanen S* (joint last authors). 2020.  Testing for context-dependent effects of maternal thyroid hormones on offspring survival and physiology: and experimental approach manipulating temperature in a wild bird species. Scientific Reports.

[46] Muola A, Fuchs B, Laihonen M, Rainio K, Heikkonen L, Ruuskanen S, Saikkonen K, Helander M. 2021. Risk in the circular food economy: Glyphosate-based herbicide residues in manure fertilizers decrease crop yield. for Scence of the Total Environment, 750, 141422.

Featured e.g. in  EurekAlert, Science Daily, Turun sanomat, Aamuset

Glyphosate residue in manure fertilizer decrease strawberry and meadow fescue growth — ScienceDaily

(Award winning) Carey Gillam (U.S. Right to know):

Glyphosate in chicken poop used as fertilizer is hurting food production, researchers say


[45] Ruuskanen S, Rainio M, Gomez-Gallogo C, Selenius O, Salminen S, Collado MC, Saikkonen K, Saloniemi I, Helander M. 2020. Glyphosate-based herbicides influence gut microbiome, reproductive hormones and oxidative status: a long-term experiment in an avian model. Environmental Pollution, 266, 115108.

Ruuskanen S, Helander M. 2020. Glyfosaatin vaikutuksia tutkittiin linnuilla. Suomen Siipikarjalehti 1/2020.

[44] Poranen O, Ruuskanen S*. 2021 Exploratory behaviour is not associated with metabolism in the Pied Flycatcher. Ibis 163, 247-252. 

[43] Ruuskanen S, Rainio M, Uusitalo M, Saikkonen K, Helander M.2020. Effects of parental exposure to glyphosate-based herbicides on embryonic development and oxidative status: a long-term experiment in a bird model. Scientific Reports 10, 6349.

[42] Sarraude T, Hsu BY, Groothuis AGG, Ruuskanen S*.2020 Manipulation of prenatal thyroid hormones does not affect growth or physiology in nestling pied flycatchers. Physiological and Biochemical Zoology,93(4):255-266, doi: 10.1086/709030.

[41] Ruuskanen S, Hsu BY, Nord A. 2021. Endocrinology of thermoregulation of birds in a changing climate. Molecular and Cellular Endocrinology, 519, 111088. EcoEvoArXiv DOI: 10.32942/

[40] Ruuskanen S, Rainio, M., Kuosmanen, V., Laihonen, M., Saikkonen, K., Saloniemi, I., Helander, M. 2019. Female preference and adverse developmental effects  of glyphosate-based herbicides on ecologically relevant traits in Japanese quails.  Environmental Science and Technology, 54, 2, 1128–1135

Featured in Helsingin Sanomat, Kemia-lehti, Siipikarjalehti, Teollisuusliitto, YLE

[39] Stier A, Bize P, Hsu B-Y, Ruuskanen S*. 2019. Plastic but repeatable: rapid adjustments of mitochondrial function and density during reproduction in a wild bird species. Biology Letters 15(11):20190536. doi: 10.1098/rsbl.2019.0536

Featured in Journal of Experimental Biology

[38] Hsu BY, Verhagen I, Gienapp P, Darras VM, Visser ME, Ruuskanen S*. 2019. Between-and within-individual variation of maternal thyroid hormone deposition in wild great tits (Parus major). American Naturalist 194, E96-E108.

[37] Morosinotto C, Thomson RL, Korpimäki E, Mateo R, Ruuskanen S*. 2019. Maternal food supplementation and perceived predation risk affect egg composition and eggshell traits but not offspring condition. Journal of Experimental Biology 222, jeb201954.

[36] Ruuskanen S, Espín S, Sánchez-Virosta P, Sarraude T, Hsu B-Y, Pajunen P, Costa RA, Eens M, Hargitai R, Török J, Eeva T. 2019. Transgenerational endocrine disruption: does elemental pollution affect egg or nestling thyroid hormone levels in a wild songbird? Environmental Pollution 247: 725-735

[35]Hsu B-Y, Doligez B, Gustafsson L, Ruuskanen S*. 2018. Transient growth‐enhancing effects of elevated maternal thyroid hormones at no apparent oxidative cost during early postnatal period. Journal of Avian Biology, DOI:  10.1111/jav.01919

[34] Ruuskanen S, Hsu B-Y, Heinonen A, Darras VM, Vainio M, Sarraude T, Rokka A. 2018. A new method for measuring thyroid hormones using nano-LC-MS/MS. Journal of Chromatography B 1093-94:24-30.

[33] Ruuskanen S & Hsu B-Y. 2018. Maternal thyroid hormones – an unexplored mechanism underlying maternal effects in an ecological framework. Physiological and Biochemical Zoology, 91(3): 904-916.

[32] Morosinotto C,  Rainio M, Ruuskanen S, Korpimäki E. 2018. Antioxidant Enzyme Activities Vary with Predation Risk and Environmental Conditions in Free-Living Passerine Birds. Physiological and Biochemical Zoology, 91(3): 837-848

[31] Ruuskanen S, Groothuis TGG; Baugh A, Schaper S, deVries B, van Oers K. 2017. Maternal egg hormones in the mating context: effect of pair personality. Functional Ecology 32:439-449.

[30] Rainio M, Ruuskanen S, Eeva T. 2017.  Spatio-temporal variation in the body condition of female pied flycatcher (Ficedula hypoleuca) in a polluted environment. Urban Ecosystems, 20:1035-1043.

[29] Ruuskanen S, Morosinotto RL, Thomson RL, Pradeep Ratnayake C, Korpimaki E. 2017 Food supplementation, but not predation risk, alters female antioxidant status during breeding. Behavioral Ecology and Sociobiology, 71:4. DOI: 10.1007/s00265-017-2299-z

[28] Ruuskanen S, Darras V, Visser ME, Groothuis TGG. 2016. Effects of experimentally manipulated yolk thyroid hormone levels on offspring development in a wild bird. Hormones and Behavior 81: 38-44.

[27] Ruuskanen S, Groothuis TGG, Gienapp P, Schaper SV, Darras VM, Pereira C, deVries B, Visser ME. 2016. Heritable variation in maternally-derived yolk androgens, thyroid hormones and immune factors. Heredity 117: 184-190.

[26] Ruuskanen S, Groothuis TGG, Schaper, SV, Darras VM, deVries B, Visser ME. 2016. Temperature-induced variation in yolk androgen and thyroid hormone levels in avian eggs. General and Comparative Endocrinology 235: 29-37.

[25] Lessells CM, Ruuskanen S, Schwabl H. 2016. Yolk hormones in great tits: multiple correlates of dihydrotestosterone. Behavioral Ecology and Sociobiology 70: 843-856.

[24] Morosinotto C, Thomson RL, Ruuskanen S, Korpimäki E, Lehikoinen E, Möstl E, Laaksonen T. 2016. Maternal deposition of androgens in eggs is affected by food availability but not by predation risk. Journal of Avian Biology 47(5).

[23] Ruiz, S, Espin S, Rainio M, Ruuskanen S, Salminen J-P, Lilley T, Eeva T. 2015. Effects of dietary lead exposure on vitamin levels in great tit nestlings – an experimental manipulation. Journal of Environmental Research 213: 688-697.

[22]Ruuskanen S. 2015. Hormonally–mediated maternal effects in birds: lessons from the flycatcher model system. General and Comparative Endocrinology 244: 283-293.

[21]Ruuskanen S, Darras V, DeVries B, Visser ME, Groothuis TGG. 2015. Experimental manipulation of food availability leads to short-term intra-clutch adjustment in egg mass but not in yolk androgen or thyroid hormones. Journal of Avian Biology 47: 36-46.

[20] Koski T-M, Laaksonen T, Mäntylä E, Ruuskanen S, Li T, Girón-Calva PS, Huttunen L, Blande JD, Holopainen JK, Klemola T. 2015. Do insectivorous birds use volatile organic compounds from plants as foraging cues? Three experimental tests. Ethology 121: 1131-1144.

[19] Ruuskanen S, Eeva T, Kotitalo P, Stauffer J, Rainio M. 2015. No delayed behavioral and phenotypic responses to experimental early-life lead exposure in great tits (Parus major). Environmental Science and Pollution Research 22: 2610-2621.

[18] Rainio M, Eeva T, Lilley T, Stauffer J, Ruuskanen S*. 2014. Effects of early-life lead exposure on oxidative status and phagocytosis activity in great tits (Parus major). Comparative Biochemistry and Physiology 167:24-34

[17] Eeva T, Rainio M, Berglund Å, Stauffer J, Stöwe M, Ruuskanen S*. 2014. Experimental manipulation of dietary lead levels in great tit nestlings –limited effects on growth, physiology and survival. Ecotoxicology 23:914-928

[16] Ratnayake CP, Morosinotto C, Ruuskanen S, Villers A, Thomson RL. 2014. Passive Integrated Transponders (PIT) on a small migratory passerine bird: absence of deleterious short and long-term effects. Ornis Fennica 91:244-255.

[15] Ruuskanen S, Laaksonen T, Morales J, Moreno J, Mateo R, Belskii E, Bushuev A.,Järvinen, A., Kerimov, A., Krams, A., Morosinotto, C., Mänd, R., Orell, M, Qvarnström A., Slater, F., Tilgar, V., Visser, M.E., Winkel, W., Zang, H. & Eeva, T. 2013. Large-scale geographical variation in eggshell metal and calcium content in a passerine bird (Ficedula hypoleuca). Environmental Science and Pollution Research 21: 3304-3317.

[14] Morosinotto C, Ruuskanen S, Thomson RL, Siitari H, Korpimäki E., Laaksonen T. 2013. Predation risk affects the levels of maternal immune factors in avian eggs. Journal of Avian Biology 44: 427-436.

[13] Ruuskanen, S, Laaksonen T. 2013. Sex-specific effects of yolk androgens on begging behavior and digestion in pied flycatchers. Journal of Avian Biology 44: 331-338

[12] Ruuskanen S, Lehikoinen E, Nikinmaa M, Siitari H, Waser W, Laaksonen, T. 2013. Long-lasting effects of yolk androgens on phenotype in the pied flycatcher (Ficedula hypoleuca). Behavioral Ecology and Sociobiolology 67:361-372

[11] Ruuskanen S, Doligez B, Gustafsson L, Laaksonen T. 2012. Long-term effects of yolk androgens on phenotype and parental feeding behavior in a wild passerine. Behavioral Ecology and Sociobiology 66: 1201-1201.

[10] Leskinen P, Laaksonen T, Ruuskanen S, Primmer C, Leder E. 2012. The proteomics of feather development in pied flycatchers (Ficedula hypoleuca) with different plumage coloration. Molecular Ecology 21:572-77

[9] Morales J, Ruuskanen S, Laaksonen T, Eeva T, Mateo R, Belskii E, Ivankina, EV, Järvinen A, Kerimov A, Korpimäki E, Krams I, Mänd R, Morosinotto C, Orell M, Qvarnström A, Siitari H, Slater FM, Tilgar V, Visser ME, Winkel W, Zang H, Moreno J. 2012. Variation in eggshell traits between geographically distant populations of pied flycatchers Ficedula hypoleuca. Journal of Avian Biology 44:112-120.

[8] Ruuskanen S, Doligez B, Pitala N, Gustafsson L, Laaksonen T. 2012. Long-term fitness consequences of high yolk androgen levels: sons pay the costs. Functional Ecology 26, 884–894

[7] Ruuskanen S, Siitari H, Eeva T, Belskii E, Järvinen A, Kerimov A, Krams I, Moreno J, Morosinotto C, Mänd R, Möstl E, Orell M, Qvarnström A, Salminen J-P, Slater F, Tilgar V, Visser ME, Winkel W, Zang H, Laaksonen T. 2011 .Geographical variation in egg mass and egg content in a passerine bird. PloS ONE 6, e25360.

[6] Merilaita S, Vallin A, Kodandaramaiah U, Dimitrova M, Ruuskanen S, Laaksonen T. 2011. Is more better? – number of eyespots and their intimidating effect on naive predators in the peacock butterfly. Behavioral Ecology 22, 1326-1331.

[5] Ruuskanen S, Helle S, Ahola M, Adamczyck F, Möstl E, Laaksonen T. 2011. Digit ratios have poor indicator value in a wild bird population. Behavioral Ecology and Sociobiolology 65, 983-994.

[4] Eeva T, Ruuskanen S, Salminen J-P, Belskii E, Järvinen A, Kerimov A, Korpimäki E, Krams I, Moreno J, Morosinotto C, Mänd R, Orell M, Qvarnström A, Siitari H, Slater FM, Tilgar V, Visser ME, Winkel W, Zang H, Laaksonen T. 2010. Geographical trends in the yolk carotenoid composition of the pied flycatcher (Ficedula hypoleuca). Oecologia 165, 277-287.

[3] Ruuskanen S, Laaksonen T. 2010. Yolk hormones have sex-specific long-term effects on behavior in the pied flycatcher (Ficedula hypoleuca). Hormones and Behavior 57, 119-127

[2] Pitala N, Ruuskanen, S., Laaksonen, T., Doligez, B., Tschirren, B. & Gustafsson, L. 2009. The effects of experimentally manipulated yolk androgens on growth and immune function of male and female nestling collared flycatchers Ficedula albicollis. Journal of Avian Biology 40, 225-230.

[1] Ruuskanen S., Doligez, B., Tschirren, B., Pitala, N., Gustafsson, L., Groothuis, T.G.G. & Laaksonen, T. 2009. Yolk androgens do not appear to mediate sexual conflict over parental investment. Hormones and Behavior 55, 514-519.

Ruuskanen S., Rainio, M., Gomez-Gallego, C., Selenius O., Salminen, S., Collado, MC., Kuosmanen V., Saikkonen, K., Saloniemi I., Helander M. 2019. Effects of glyphosate-based herbicides on non-target avian taxa: development, reproduction and fecal microbiome. SETAC conference proceedings, 25.9.2019