Muola, A., Birge, T., Helander, M., Mathew, SA., Harazinova, V., Saikkonen, K., Fuchs, B., 2023. Endophytic Beauveria bassiana induces biosynthesis of flavonoids in oilseed rape following both seed inoculation and natural colonization Pest Management Science in press.

Helander, M., Jeevannavar, A., Kaakinen, K., Mathew, SA., Saikkonen, K., Fuchs, B., Puigbò, P., Loukola, OJ., Tamminen, M., 2023. Glyphosate and a glyphosate-based herbicide affect bumblebee gut microbiota FEMS Microbiology Ecology, fiad065

Fuchs B., Saikkonen K., Damerau A., Yang B., Helander M., 2023. Herbicide residues in soil decrease microbe-mediated plant protection. Plant Biology 25 (4), 571-578

Rainio M.J., Margus A., Tikka S., Helander M., Lindström L., 2023. The effects of short-term glyphosate-based herbicide exposure on insect gene expression profiles. Journal of Insect Physiology 146: 104503

Mathews, S.A., Fuchs, B., Saikkonen, K., Helander, M., Puigbo, P., Nissinen, R., Muola, A. 2023. Glyphosate-based herbicide use affects individual microbial taxa in strawberry endosphere but not the microbial community composition. Journal of Applied Microbiology lxad006

Mathew, S.A., Helander, M., Saikkonen, K., Vankova, R., Dobrev, P., Dirihan, S., Fuchs B. 2023. Epichloë endophytes shape the foliar endophytic fungal microbiome and alter the auxin and salicylic acid phytohormone levels in two meadow fescue cultivars. Journal of Fungi 9 (1), 90

Kalske, A., Saikkonen, K., Helander, M. 2023. Endophytic Fungus Negatively Affects Salt Tolerance of Tall Fescue Journal of Fungi 9 (1), 14

Keronen S., Helander, M., Saikkonen K., Fuchs B. 2023. Management practice and soil properties affect plant productivity and root biomass in endophyte-symbiotic and endophyte-free meadow fescue grasses Journal of Sustainable agriculture and Environment. 2: 16-25

Ruuskanen, S., Fuchs, B., Nissinen, R., Puigbò, P., Rainio, M., Saikkonen, K., Helander, M., 2023. Ecosystem consequences of herbicides: the role of microbiome. Trends in Ecology & Evolution 0.

Helander, M., Lehtonen, T.K., Saikkonen, K., Despains, L., Nyckees, D., Antinoja, A., Solvi, C., Loukola, O.J., 2023. Field-realistic acute exposure to glyphosate-based herbicide impairs fine-color discrimination in bumblebees. Science of The Total Environment 857, 159298.


Laihonen, M., Rainio K., Birge, T., Saikkonen, K., Helander, M., Fuchs, B. 2022. Root biomass and cumulative yield increase with mowing height in Festuca pratensis irrespective of Epichloë symbiosis. Scientific Reports 12 (1), 1-10

Fuchs, B., Saikkonen, K., Helander, M., Tian, Y., Yang, B., Salminen, J-P., Muola, A. 2022 Legacy of agrochemicals in the circular food economy: Glyphosate-based herbicides introduced via manure fertilizer affect the yield and biochemistry of perennial crop plants during the following year. Chemosphere, 136366.

Ramula S., Kalske A., Saikkonen K., Helander M. 2022 Glyphosate residues in soil can modify plant resistance to herbivores through changes in leaf quality. Plant Biology

Mathew S.A., Muola A., Saikkonen, K., Saloniemi I., Helander M. & Puigbò P. 2022 Quantification of the Potential Impact of Glyphosate-Based Products on Microbiomes. , 10 Jan 2022, (179)10.3791/63109

Puigbo P., Leino L.I., Rainio M.J., Saikkonen K., Saloniemi I., Helander M., 2022. Does Glyhosate Affect the Human Microbiota? Life. 12(5), 707;

De-la-Cruz I.M., Batsleer F., Bonte D., Diller C., Hytönen T., Muola A., Osorio S., Posé D., Vandegehuchte M.L. & Stenberg J.A. 2022. Evolutionary Ecology of Plant-Arthropod Interactions in Light of the “Omics” Sciences: A Broad Guide. Front. Plant Sci. 13:808427. doi: 10.3389/fpls.2022.808427

Ruotsalainen A. L., Kauppinen M., Wäli P. R., Saikkonen K., Helander M. & Tuomi J. 2022. Dark septate endophytes: by-product mutualists and tomorrow’s mycorrhizal symbionts? Trends in Plant Science 27 (3): 247-254. doi: 10.1016 / j.tplants.2021.10.001.

Fuchs B., Laihonen M., Muola A., Saikkonen K., Dobrev P. I., Vankova R., Helander M. 2022. A glyphosate-based herbicide in soil differentially affects hormonal homeostasis and performance of non-target crop plants. Frontiers in Plant Science 12:787958.

Laihonen, M., Saikkonen, K., Helander, M., Vázquez de Aldana, B.R., Zabalgogeazcoa, I., Fuchs, B., 2022. Epichloë endophyte-promoted seed pathogen increases host grass resistance against insect herbivory. Frontiers in Microbiology 12:786619.

Schott, J., Fuchs, B., Böttcher, C., Hilker, M., 2022. Responses to larval herbivory in the phenylpropanoid pathway of Ulmus minor are boosted by prior insect egg deposition. Planta 255, 16.


Egan P. A., Muola M., Parachnowitsch A. l., Stenberg J. A. 2021. Pollinators and herbivores interactively shape selection on strawberry defense and attraction. Evolution Letters, doi: 10.1002 / evl3.262.

Koski T. M., De Jong S., Muola A., Buchvaldt Amby D., Andreasson E., Stenberg J. A. 2021. ‘Resistance mixtures’ reduce insect herbivory in strawberry (Fragaria vesca) plantings. Frontiers in Plant Science 12: 722795.

Von Cräutlein M., Helander M., Korpelainen H., Leinonen P. H., Vázquez de Aldana B. R., Young C. A., Zabalgogeazcoa I., Saikkonen K. 2021. Genetic diversity of the symbiotic fungus Epichloë festucae in naturally occurring host grass populations. Frontiers in Microbiology 12:756991.

Ramula S., Mathew S. A., Kalske A., Nissinen R., Saikkonen K. and Helander M. 2021. Glyphosate residues alter the microbiota of a perennial weed with minimal indirect impact on plant performance. Plant and Soil .

Muola, A., Scheepens, JF, Laukkanen, L, Kalske, A., Mutikainen, P. and Leimu, R. 2021. Strong gene flow explains lack of mating system variation in the perennial herb, Vincetoxicum hirundinaria , in a fragmented landscape. Nordic Journal of Botany, 39:.

Fuchs, B., Saikkonen, K., Helander, M. 2021. Glyphosate-modulated biosynthesis driving plant defense and species interactions. Trends in Plant Science 26: 312-323 doi:

Rainio, M., Ruuskanen, S., Helander, M., Saikkonen, K., Saloniemi, I., Puigbò, P. 2021. Adaptation of Bacteria to Glyphosate: a microevolutionary perspective of the enzyme 5-enolpyruvylshikimate-3-phosphate synthase. Environmental Microbiology Reports, doi:

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. Science of the Total Environment, doi:


Leino, L., Tall, T., Helander, M., Saloniemi, I., Saikkonen, K., Ruuskanen, S., Puigbò, P. 2020: Classification of the Glyphosate target enzyme (5-enolpyruvylshikimate-3-phosphate synthase) for assessing the sensitivity of organisms to the herbicide. Journal of Hazardous Materials, doi:

Ruuskanen, S., Rainio, M., Gomez-Gallego, C., Selenius, O., Salminen, S., Carmen Collado, M. Saikkonen, K., Saloniemi, I. & Helander, M. 2020: Glyphosate- based herbicides influence antioxidants, reproductive hormones and gut microbiome but not Reproduction: A long-term experiment in an avian model. Environmental Pollution, DOI: 10.1016 / j.envpol.2020.115108

Rainio, M., Margus, A., Virtanen, V., Lindström, L., Salminen, JP., Saikkonen, K., Helander, M. 2020: Glyphosate-based herbicides have soil-mediated effects on potato glycoalkaloids and oxidative status of a potato Pest. Chemosphere, DOI: 10.1016 / j.chemosphere.2020.127254

Vázquez de Aldana, BR, Leinonen, P., Zabalgogeazcoa, I., Helander, M. & Saikkonen, K. 2020: Variation and plasticity in Epichloë alkaloid content of Festuca rubra across Europe. Fungal Ecology 47: 100942

Ruuskanen S., Rainio M., Uusitalo M., Saikkonen, K., Saloniemi, I. & 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

Weber D, Egan PA, Muola A, Ericson LE & Stenberg JA. 2020: Plant resistance does not compromise parasitoid-based biocontrol of a strawberry pest. Scientific Reports 10: 5899

Aikio S, Ramula S, Muola A & von Numers M. 2020: Island properties dominate species traits in determining plant colonizations in an archipelago system. Ecography 43: 1-11

Gómez-Gallego, C., Rainio, M., Collado, M., Mantziari, A., Salminen, S., Saikkonen, K., Helander, M. 2020: Glyphosate-based herbicide affects the composition of microbes associated with Colorado potato beetle ( Leptinotarsa ​​decemlineata ). FEMS Microbiology Letters, fnaa050. DOI: 10.1093 / femsle / fnaa050

Fuchs, B., Kuhnert, E., Krauss, J. 2020: Contrasting Effects of Grass – Endophyte Chemotypes on a Tri-Trophic Cascade. Journal of Chemical Ecology. DOI: 10.1007 / s10886-020-01163-9

Saikkonen, K., Nissinen, R. & Helander, M. 2020: Towards comprehensive plant microbiome research. Frontiers in Ecology and Evolution. DOI: 10.3389 / fevo.2020.00061

Bao, G., Song, M., Wang, Y. & Saikkonen, K. & Li, C. 2020: Does Epichloë endophyte enhance host tolerance to root hemiparasite? Microbial Ecology DOI: 10.1007 / s00248-020-01496-8

Ruuskanen, S., Rainio, M., Kuosmanen, V., Laihonen, M., Saikkonen, K., Saloniemi, I. & Helander, M. 2020: Female preference and adverse developmental effects of Glyphosate-based herbicide on ecologically relevant traits in Japanese quails. Environmental Science & Technology. DOI: 10.1021 / acs.est.9b07331


Bao, G., Song, M., Wang, Y., Saikkonen, K. & Wang, H. 2019: Interactive effect of Epichloë fungal and host Origins on seed germination of Achnatherum inebrians . Symbiosis 79: 49-58

Hagner, M., Mikola, J., Saloniemi, I., Saikkonen, K. & Helander, M. 2019: Effects of a glyphosate-based herbicide on soil animal trophic groups and associated ecosystem functioning in a northern agricultural field. Scientific Reports 9: 8540, DOI: org / 10.1038 / s41598-019-44988-5

Helander, M., Pauna, A., Saikkonen, K. & Saloniemi, I. 2019. Glyphosate residues in soil affect crop plant germination and growth. Scientific Reports 9:19653 DOI: org/10.1038/s41598-019-56195-3

Kauppinen, M., Helander, M., Anttila, N., Saloniemi, I., Saikkonen, K. 2019: Epichloë endophyte effects on leaf blotch pathogen (Rhynchosporium sp.) of tall fescue vary among grass origin and environmental conditions. Plant Ecology and Diversity. DOI: 10.1080/17550874.2019.1613451

Koski, T.-M., Saikkonen, K., Klemola, T. & Helander, M. 2019: Foraging preferences of barnacle geese on endophytic tall- and red fescues. Human–Wildlife Interactions 13(2): 331-343

Leinonen, P.H., Helander, M., Vázquez-de-Aldana, B.R., Zabalgogeazcoa, I., Saikkonen, K. 2019: Local adaptation in natural European host grass populations with asymmetric symbiosis. PLos One 14(4): e0215510, DOI: org/10.1371/journal.pone.0215510

Margus, A., Rainio, M. & Lindström, L. 2019: Can Indirect Herbicide Exposure Modify the Response of the Colorado Potato Beetle to an Organophosphate Insecticide? Journal of Economic Entomology, toz115,

Nissinen, R., Helander, M., Kumar, M. & Saikkonen, K. 2019: Heritable Epichloë symbiosis shapes fungal but not bacterial communities of plant leaves. Scientific Reports 9: 5253, DOI: org/10.1038/s41598-019-41603-5

Rainio, MJ, Margus, A., Lehmann, P., Helander, M. & Lindström, L. 2019: Effects of a Glyphosate-based herbicide on Survival and oxidative status of a non-target herbivore, the Colorado potato beetle (Leptinotarsa decemlineata). Comparative Biochemistry and Physiology, Part C 215: 47-55, DOI: 10.1016 / j.cbpc.2018.09.005

Saikkonen, K., Dirihan, S., Väre, H., Saloniemi, I., von Cräutlein, M., Leinonen, P. & Helander, M. 2019. Phenotypic and Genetic Variation in Natural Populations of Festuca rubra  sl sensu lato in Europe. Plant Ecology and Diversity DOI: org / 10.1080 / 17550874.2019.1654551

Saravesi K, Markkola A, Taulavuori E, Syvänperä I, Suominen O, Suokas M, Saikkonen K, Taulavuori K, 2019. Impacts of experimental warming and northern light climate on growth and root fungal communities of Scots pine populations. Fungal Ecology. doi:

von Cräutlein, M., Leinonen, PH, Korpelainen, H., Helander, M., Väre, H. & Saikkonen, K. 2019: Post-glacial Colonization history reflects in the Genetic structure of natural populations of Festuca rubra in Europe. Ecology and Evolution DOI: 10.1002 / ece3.4997


Compant, S., Saikkonen, K., Mitter, B., Campisano, A. & Mercado-Blanco, J. 2016: Editorial special issue: Soil, Plants and Endophytes. Plant and Soil 405: 1-11. DOI: 10.1007 / s11104-016-2927-9

Dirihan, S., Helander, M., Väre, H., Gundel, PE, Garibaldi, LA, Irisarri, JGN, Saloniemi, I. & Saikkonen, K. 2016: Geographic variation in Festuca rubra L. ploidy levels and systemic fungal endophyte frequencies. PLOS ONE DOI: 10.1371 / journal.pone.0166264

Fält-Nardmann, J., Klemola, T., Roth, M., Ruohomäki, K. & Saikkonen, K. 2016: Northern geometrid forest pests (Lepidoptera: Geometridae) Hatch at lower temperatures than their southern conspecifics: implications of climate change . Europ. J. Entomol. 113: 337-343.

Gundel, PE, Helander, M., Garibaldi, LA, Vázquez-de-Aldana, BR, Zabalgogeazcoa, I. & Saikkonen, K. 2016: Role of foliar fungal endophytes in litter decomposition among species and population Origins. Fungal Ecology 21: 50-56.

Helander, M., Phillips, T., Faeth, SH, Bush, LP, McCulley, R., Saloniemi, I. & Saikkonen, K. 2016: Alkaloid quantities in endophyte infected tall fescue are affected by the plant-fungus combination and environment. Journal of Chemical Ecology 42: 118-126.

Hume, DE, Geraldine, DR, Gibert, A., Helander, M., Mirlohi, A. & Sabzalian, MR 2016: Epichloë fungal endophytes for grassland ecosystems. (Lichtfouse, E., ed.) Sustainable Agriculture Reviews 19: 233-305.

Kauppinen, M., Saikkonen, K, Helander, M., Pirttilä, A.M. & Wäli, P.R. 2016: Epichloë grass endophytes in sustainable agriculture in changing climate. Nature Plants 2: 15224. doi:10.1038/nplants.2015.224.

Mikola, J., Helander, M. & Saikkonen, K. 2016. No effects of Epichloë endophyte infection on nitrogen cycling in meadow fescue (Scherodonus pratensis) grassland. Plant and Soil 405: 257-264. DOI 10.1007/s11103-015-2711-2

Ovčarenko, I., Lindström, L., Saikkonen, K., Jauhiainen, L., Kaseva, J. & Vänninen, I.  2016: Preconditioning of the generalist herbivore Trialeurodes vaporariorum to greenhouse monocultures and its subsequent performance on wild polycultures. Entomol. Experim. Applic. 159: 1-16.

Saikkonen, K. Young, CA, Helander, M. & Schardl, CL 2016: Endophytic Epichloë species and their grass hosts: from Evolution to applications. Plant Molecular Biology 90: 665-675. DOI: 10.1007 / s11103-015-0399-6

Saikkonen, K., Phillips, TD, Faeth, SH, McCulley, RL, Saloniemi, I. & Helander, M. 2016: Performance of endophyte infected tall fescue in Europe and North America. PLoS ONE 11: e0157382

van Overbeek, LS & Saikkonen, K. (2016): Impact of bacterial-fungal interactions on the colonization of the endosphere. Trends Plant Sci. 21: 230-242. DOI: org / 10.1016 / j.tplants.2016.01.003


Gundel, PE, Dirihan, S., Helander, M., Zabalgogeazcoa, I., Väre, H. & Saikkonen, K. 2014: Systemic fungal endophytes and ploidy levels in Festuca vivipara populations in the North European Islands. Plant Systematics and Evolution 300: 1683-1691. DOI: 10.1007 / s00606-014-0994-z

Gundel, PE, Garibaldi, LA, Wäli, PR, Helander, M., Dirihan, S. & Saikkonen, K. 2014: Fungal endophyte mediated occurrence of seminiferous and pseudoviviparous panicles in Festuca rubra . Fungal Diversity 66: 69-76. DOI: 10.1007 / s13225-014-0290-9

Huitu, O., Forbes, K., Helander, M., Julkunen-Tiitto, R., Lambin, X., Saikkonen, K., Stuart, P., Sulkama, S. & Hartley, SE 2014: Silicon, endophytes and secondary metabolites as grass defenses against mammalian herbivores. Frontiers in Plant Science 5: e478. DOI: 10.3389 / fpls.2014.00478

Li, T., Blande, JD, Gundel, PE, Helander, M. & Saikkonen, K. 2014: Epichloë endophytes alter inducible indirect defenses in host grasses. PLOS ONE  9: e10133. DOI: 10.1371 / journal.pone.0101331

Ovčarenko, I., Lindström, L., Saikkonen, K. & Vänninen, I. 2014: Variation in mortality among populations is higher for pymetrozine than for imidacloprid and spiromesifen in Trialeurodes vaporariorum in Greenhouses in Finland. Pest Manag. Sci. 70: 1524–1530.

von Cräutlein, M., Korpelainen, H., Helander, M., Öhberg, A. & Saikkonen, K. 2014: Development and characterization of nuclear microsatellite markers in the endophytic fungus   Epichloë festucae (Clavipitaceae). Applications in Plant Sciences 2: 1400093. DOI: org / 10.3732 / apps.1400093

von Cräutlein, M., Korpelainen, H., Helander, M., Väre, H. & Saikkonen, K. 2014: Development and characterization of chloroplast microsatellite markers in fine-leaved fescue, Festuca rubra (Poaceae). Applications in Plant Sciences 2: 1400094. DOI: org / 10.3732 / apps.1400094


Dirihan, S., Terho, P., Helander, M. & Saikkonen, K. 2013: Efficient analysis of ploidy levels in plant Evolutionary ecology. Caryology: International Journal of Cytology, Cytosystematics and Cytogenetics 66: 251-256.

Gundel, PE, Garibaldi, LA, Helander, M. & Saikkonen, K. 2013: Symbiotic interactions as drivers of trade-offs in plants: effects of fungal endophytes on tall fescue. Fungal Diversity 60: 5–14.

Gundel, PE, Helander, M., Casas, C., Hamilton, CE, Faeth, S. & Saikkonen, K. 2013: Neotyphodium fungal endophyte in tall fescue ( Schedonorus phoenix ): A comparison of three Northern European wild populations and the cultivar Kentucky-31. Fungal Diversity 60: 15–24.

Gundel, PE, Pérez, LI, Helander, M. & Saikkonen, K. 2013. Symbiotically modified organisms: nontoxic fungal endophytes in grasses. Trends in Plant Science 18: 420-427.

Helander, M., Jia, R., Huitu, O., Sieber, TN, Jia, J., Niemelä, P. & Saikkonen, K. 2013: Endophytic fungi and silica content of different Bamboo species in Giant panda diet. Symbiosis 61: 13–22.

Martín, J.A., Witzell, J., Blumenstein, K., Rozpedowska, E., Helander, M., Sieber, T. & Gil, L. 2013: Resistance to Dutch elm disease restricts xylem endophytic fungi in elms (Ulmus spp.). PLoS ONE 8: e56987. doi:10.1371/journal.pone.0056987.

Saikkonen, K., Gundel, P. & Helander, M. 2013: Chemical ecology mediated by fungal endophytes in grasses. Journal of Chemical Ecology 39:962–968.

Saikkonen, K., Ruokolainen, K., Huitu, O., Gundel, P.E., Piltti, T., Hamilton, C.E. & Helander, M. 2013: Fungal endophytes help prevent weed invasions. Agriculture, Ecosystems and Environment 165: 1– 5.

Zabalgogeazcoa, I., Gundel, PE, Helander, M. & Saikkonen, K. 2013: Non-systemic fungal endophytes in Festuca rubra plants infected by Epichloë festucae in subarctic habitats. Fungal Diversity 60: 25–32.


Ammunét, T., Kaukoranta, T., Saikkonen, K., Repo T. & Klemola, T. 2012: Invasive and Resident defoliators in a changing climate: cold Tolerance and predictions concerning extreme winter cold as a range limiting factor. Ecol. Entomol. 37: 212-220.

Gundel, PE, Hamilton, CE, Seal, CE, Helander, M., Martínez-Ghersa, MA, Ghersa, CM, Vázquez de Aldana, BR, Zabalgogeazcoa, I. & Saikkonen, K. 2012: Antioxidants in Festuca rubra L. seeds affected by the fungal symbiont Epichloë festucae . Symbiosis 58: 73-80.

Hamilton, C., Gundel, PE, Helander, M. & Saikkonen, K. 2012: Endophytic Mediation of reactive oxygen species and Antioxidant activity in plants: a review. Fungal Diversity , 54: 1-10. DOI 10.1007 / s13225-012-0158-9

Haviola, S., Neuvonen, S., Rantala, MJ, Saikkonen, K., Salminen, J.-P., Saloniemi, I., Yang, S. & T. Ruuhola 2012: Genetic and environmental factors behind foliar Chemistry of the mature mountain birch. J. Chem. Ecol. 38: 902–913.

Helander, M., Saloniemi, I. & Saikkonen, K. 2012: Glyphosate in northern ecosystems. Trends in Plant Science 17: 569-574.

Liimatainen, J., Karonen, M., Sinkkonen, J., Helander, M. & Salminen, J.-P. 2012: Characterization of phenolic compounds from inner bark of Betula pendula. Holzforschung 66: 171-181. DOI: 10.1515/HF.2011.146

Liimatainen, J., Karonen, M., Sinkkonen, J., Helander, M. & Salminen, J.-P. 2012: Phenolic compounds of the inner bark of Betula pendula: Seasonal and genetic variation and induction by wounding. Journal of Chemical  Ecology 38:1410-1418. DOI 10.1007/s10886-012-0199-2.

Saikkonen, K., Taulavuori, K., Hyvönen, T., Gundel, PE, Hamilton, CE, Vänninen, I. Nissinen, A. & Helander, M. 2012. Climate change-driven species’ range shifts filtered by photoperiodism. Nature Climate Change 2: 234-242. DOI: 10.1038 / NCLIMATE1430