Original research articles


  1. Järvinen, L., Santavirta, S., Putkinen, V., Karlsson, H.K., Seppälä, K., Sun, L., Hudson, M., Hirvonen, J., Nuutila, P., & Nummenmaa, L. (preprint). Reward responses to vicarious feeding depend on BMI. bioRxiv.
  2. Lahnakoski, J.M., Bennett, E., Nummenmaa, L., Steinert, U., Sams, M., & Svärd, S. (preprint). Embodied emotions in ancient Neo-Assyrian texts revealed by bodily mapping of emotional semantics. PsyArXiv.
  3. Seppälä, K., Putkinen, V., Harju, H., Rebelos, E., Hirvonen, J., Helin, S., Rajander, J., Karlsson, H., Saunavaara, J., Hyönä, J., & Nummenmaa, L. (preprint). Endogenous opioid system modulates proximal and distal threat signals in the human brain. bioRxiv.
  4. Putkinen, V., Seppälä, K., Harju, H., Hirvoinen, J., Karlsson, H.K., & Nummenmaa, L. (preprint). Pleasurable music activates cerebral μ-opioid receptors: A combined PET-fMRI study. bioRxiv.
  5. Noppari. T., Tuisku, J., Lukkarinen, L., Tani, P., Lindberg, N., Saure, E., Lauerma, H., Tiihonen, J., Hirvonen, J., Helin, S., Rajander, J., Salmi, J., & Nummenmaa, L. (preprint). Dopamine D2R and opioid MOR availability in autism spectrum disorder. bioRxiv
  6. Chen, J., Putkinen, V., Seppälä, K., Hirvonen, J., Ioumpa, K., Gazzola, V., Keysers, K., & Nummenmaa, L. (preprint). Endogenous opioid receptor system mediates costly altruism in the human brain. bioRxiv.
  7. Saanijoki, T., & Nummenmaa, L. (preprint). Bodily maps of exercise-induced sensations. psyArXiv.
  8. Pak, K., Tuisku, J., Karlsson, H.K., Hirvonen, J., Rebelos, E., Pekkarinen, L., Sun, L., Latva-Rasku, A., Helin, S., Rajander, J., Karukivi, M., Nuutila, P., & Nummenmaa, L. (preprint). Anorexia nervosa is associated with higher brain mu-opioid receptor availability. medRxiv.
  9. Pak, K.,  Malén, T.,  Santavirta, S., Shin, S.,  Nam, H.Y., De Maeyer, S., & Nummenmaa, L. (preprint). Glucose metabolism and radiodensity of abdominal adipose tissue: A 5-year longitudinal study in a large PET cohort. bioRxiv.
  10. Romero, R.R., Silvestrin, M., Glerean, E., Nummenmaa, L., & Boggio, P.S (preprint). Dehumanisation and distinct bodily emotions towards outgroup members. Human Emotion Systems Labtoratory Preprints. 
  11. Santavirta, S., Malen, T., Erdemli, A., &, Nummenmaa, L. (preprint). Taxonomy of human social perception. bioRxiv.
  12. Aksiuto, A., Lahnakoski, J.M., Kauttonen, J., Saarimäki, H., Volynets, S., Nummenmaa, L., Sams, M., & Jääskeläinen, I.P. (preprint). Psychopathic traits predict neural responses to emotional movies in the general population. bioRxiv.
  13. Lukkarinen, L., Tuisku, J., Sun, L., Helin, S., Karlsson, H.K., Venetjoki, N., Salomaa, M., Rautio, P., Hirvonen, J. Lauerma, H., Tiihonen, J., & Nummenmaa, L. (preprint). Aberrant type 2 dopamine receptor availability in criminal psychopathy. bioRxiv.
  14. Saarimäki, H., Nummenmaa, L., Volynets, S., Santavirta, S., Aksiuto, A., Sams, M., Jääskeläinen, I.P., & Lahnakoski, J. (preprint). Cerebral topographies of perceived and felt emotions. bioRxiv.
  15. Oksama, L., Heikkilä, T., Nummenmaa, L., Sams, M., & Hyönä. J. (preprint). Auditory tracking of multiple naturalistic moving objects. PsyArXiv.


  1. Nummenmaa, L. (in press). Psychology and neurobiology of horror movies. Projections: The Journal for Movies and Mind. 
  2. Latva-Rasku, Al. Rebelos, E., Bhomwik, A., Tuisku, J., Keskinen, H., Laurila, S., Lahesmaa-Hatting, M., Aarnio, R., Kirjavainen, A.K., Koffert, J., Heurling, K., Nummenmaa, L., Oscarsson, J., & Nuutila, P. (in press). SGLT2 Inhibitor Dapagliflozin Increases Skeletal Muscle and Brain Fatty Acid Uptake in People With Type 2 Diabetes – A Randomized, Double-Blind, Placebo-Controlled, Positron Emission Tomography Study. Diabetes Care. 
  3. Sun, L., Malen, T., Tuisku, J., Kaasinen, V., Hietala, J.A.m Rinne, J., Nuutila, P., & Nummenmaa, L. (2024). Seasonal variation in D2/3 dopamine receptor availability in the human brain. European Journal of Nuclear Medicine and Molecular Imaging. 
  4. Putkinen, V., Zhou, X., Gan, X., Yang, L., Becker, B., Sams, M., & Nummenmaa, L. (2024). Bodily maps of musical sensations across cultures. Proceedings of the National Academy of Sciences of the United States of America, 121, e2308859121. Data and code may be downloaded from OSF
  5. Niemi, K.J., Huovinen, A., Jaakkola, E., Glerean, E., Nummenmaa, L., & Joutsa, J. (2024). Bodily maps of symptoms and emotions in Parkinson’s disease. Movement Disorders.
  6. Malén, T., Santavirta, S., De Maeyer, S., Tuisku, J., Kaasinen, V., Kankare, T., Isojärvi, J., Rinne, J., Hietala, J., Nuutila, P., & Nummenmaa, L. (2024).  Alterations in type 2 dopamine receptors across neuropsychiatric conditions: A large-scale PET cohort. NeuroImage: Clinical.
  7. Bucci, M., Rebelos, E., Oikonen, V., Rinne, J., Nummenmaa, L., Iozzo, P., & Nuutila, P. (2024). Kinetic Modelling of Brain [18-F]FDG PET time activity curves with  Input Function Recovery (IR) method. Metabolites. 
  8. Sun, R., Balabanova, A., Bajada, C, …, Nummenmaa, L., … & Sauter, D. (2024). Emotional experiences and psychological wellbeing in 51 countries during the COVID- 19 pandemic. Emotion, 24, 397–411.


  1. Rebelos, E., Latva-Rasku, A., Koskensalo, K., Pekkarinen, L., Saukko, E., ihalainen, J., Honka, M-J., Tuisku, J., Bucci, M., Laurila, S., Rajander, J., Salminen, P., Nummenmaa, L., Jansen, J.F.A., Ferrannini, E., & Nuutila, P. (in press). Insulin-stimulated brain glucose uptake correlates with brain metabolites in morbid obesity: a combined neuroimaging study. Journal of Cerebral Blood Flow and Metabolism.
  2. Ojala, J., Suvilehto, J.T., Nummenmaa, L., & Kalso, E. (2023). Bodily maps of emotions and nociceptive, tactile and hedonic sensitivity in healthy controls and patients experiencing chronic pain. Pain.
  3. Jern, P., Chen, J., Tuisku, J., Saanijoki, T., Hirvonen, J., Lukkarinen, L., Manninen, S., Helin, S., Putkinen, V., & Nummenmaa, L. (2023). Endogenous opioid release after orgasm in man: A combined PET/functional MRI Study  Journal of Nuclear Medicine.
  4. Nummenmaa, L., Malen, T., Nazari-Farsani, S., Seppälä. K., Sun, L., Karlsson, H.K., Hudson, M., Hirvonen, J., Sams, M., Scott, S.K., Putkinen, V. (2023) Decoding brain basis of laughter and crying in natural scenes. NeuroImage.
  5. Santavirta, S., Karjalainen, T., Nazari-Farsani, S., Hudson, M., Putkinen, V., Seppälä, K., Sun, L., Glerean, E., Hirvonen, J., Karlsson, H.K., & Nummenmaa, L. (2023). Functional organization of social perception networks in the human brain. NeuroImage.
  6. Bucci, M., Iozzo, P., Merisaari, H., Huovinen, V., Lipponen, H., Räikkönen, K., Parkkola, R., Salonen, M., Sandboge, S., Eriksson, J.G., Nummenmaa, L., & Nuutila, P. (2023). Resistance training increases white matter density in frail elderly women. Journal of Clinical Medicine.
  7. Pekkarinen, L., Kantonen, T., Oikonen, V., Haaparanta-Solin, M., Aarnio, R., Dickens, A.M., , von Eyken, A., Latva-Rasku, A., Dadson, P., Kirjavainen, A.K., Helin, S., Rajander, J., Kalliokoski, K., Rönnemaa, T., Nummenmaa, L., & Nuutila, P. (2023). Lower CB1 receptor density in abdominal adipose tissue in young men with overweight. Obesity.
  8. Schaper, F.L.W.V.J., Nordberg, J., Cohen, A.L., …, Nummenmaa, L.,…, & Fox, M.D. (2023). Lesion-related epilepsy maps to a common brain network. JAMA Neurology, 80, 891–902.
  9. U-Din, M., Rebelos, E., Saari, T., Niemi, T., Eskola, O., Fromme, T., Rajander, J., Taittonen, M., Klingespor, M., Nuutila, P., Nummenmaa, L., & Virtanen, K.A. (2023). Thermogenic capacity of human supraclavicular brown fat and cold-stimulated brain glucose metabolism. Metabolites.
  10. Nordberg, J., Schaper, F.L., Bucci, M., Nummenmaa, L., & Joutsa, J. (2023). Brain lesion localizations associated with secondary seizure generalization in tumors and strokes. Human Brain Mapping, 44,3136-3146.
  11. Hudson, M., Santavirta, S., Putkinen, V., Seppälä, K., Karjalainen, T., Karlsson, H.K., Hirvonen, J., & Nummenmaa, L. (2023). Neural responses to biological motion distinguish autistic and schizotypal traits in the general population. Social Cognitive and Affective Neuroscience.
  12. Nummenmaa, L. & Hari, R. (2023). Bodily feelings and aesthetic experience of art. Cognition & Emotion, 37, 515-528.
  13. Sun, L., Laurila, S., Lahesmaa, M., Rebelos, E., Virtanen, K.A., Schnabl, K., Klingenspor, M., Nummenmaa, L., & Nuutila, P. (2023). Secretin modulates appetite via brown adipose tissue – brain axis. European Journal of Nuclear Medicine and Molecular Imaging, 50, 1597-1606.
  14. Putkinen, V. , Nazari-Farsani, S., Karjalainen, T., Santavirta, S., Hudson, M., Seppälä, K., Sun, L., Karlsson, H.K., Hirvonen, J., & Nummenmaa, L. (2023). Pattern recognition reveals sex-dependent neural substrates of sexual perception. Human Brain Mapping, 44, 2543-2556.
  15. Blain, S.D., Snodgress, M.A., Nummenmaa, L., Peterman, J.S., Glerean, E., & Park, S. (2023). Social bodies: Preliminary evidence that awareness of embodied emotions is associated with recognition of emotions in the bodily cues of others. Psychology of Consciousness: Theory, Research, and Practice.


  1. Pak, K., Kantonen, T., Pekkarinen, L., Nuutila, P., & Nummenmaa, L. (2022). Association of CNR1 gene and Cannabinoid 1 receptor protein in the human brainJournal of Neuroscience Research.
  2. Pak, K., Malen, T., Santavirta, S., Shin, S., Nam, H.-Y., De Mayer, S., & Nummenmaa, L. (2022). Brain glucose metabolism and ageing: A 5-year longitudinal study in a large PET cohort. Diabetes Care.
  3. Pekkarinen, L., Kantonen, T., Rebelos, E., Latva-Rasku, A., Dadson, P., Karjalainen, T., Bucci, M., Kalliokoski, K., Laitinen, K., Houttu, N., Kirjavainben, A.K., Rönnemaa, T., Nummenmaa, L., & Nuutila, P. (2022). Obesity risk is associated with brain glucose uptake and insulin resistance. European Journal of Endocrinology
  4. Sun, L., Aarnio, R., Atenico Herre, E., Kärnä, S., Palani, S., Virtanen, H., Liljebäck, H., Virta, J., Honkanemi, A., Oikonen, V., Han, C., Laurila, S., Bucci, M., Helin, S., Yatkin, E., Nummenmaa, L., Nuutila, P., Tang, J., & Roivainen, A. (2022). Photoperiod modulates mu-opioid receptor availability in the brown adipose tissue. European Journal of Nuclear Medicine and Molecular Imaging.
  5. Nummenmaa, L. (2022). Mapping emotions on the body. Scandinavian Journal of Pain.
  6. Noppari, T., Sun, L., Lukkarinen, L., Putkinen, V., Tani, P., Lindberg, N., Saure, E., Lauerma, H., Tiihonen, J., Venetjoki, N., Salomaa, M., Rautio, P., Hirvonen, J., Salmi, J., & Nummenmaa, L. (2022). Brain structural alterations in autism and criminal psychopathy. NeuroImage Clinical.
  7. Rebelos, E., Rissanen, E., Bucci, M., Jääskeläinen, O., Honka, M.-J., Nummenmaa, L., Moriconi, D., Laurila, S., Salminen, P., Herukka, S.-K., Singhal, T., & Nuutila, P. (2022). Circulating neurofilament is linked with obesity, renal function, and brain cerebral atrophy. Scientific Reports.
  8. Malen, T., Karjalainen, T., Isojärvi, J., Vehtari, A., Brukner, P., Putkinen, V., Kaasinen, V., Hietala, J., Nuutila, P., Rinne, J., & Nummenmaa, L. (2022). Atlas of type 2 dopamine receptors in the human brain: Age and sex dependent variability in a large PET cohortNeuroImage.
  9. Saanijoki, T., Kantonen, T., Pekkarinen, L., Kalliokoski, K., Hirvonen, J., Tuominen, L., Tuulari, J.J., Arponen, E., Nuutila, P., & Nummenmaa, L. (2022). Aerobic fitness is associated with cerebral mu-opioid receptor availability and activation in healthy humans. Medicine & Science in Sports & Exercise.
  10. Sun, L., Lukkarinen, L., Karlsson, H., Hirvonen, J., Tiihonen, J., Lauerma, H., Scott, S., & Nummenmaa, L. (2022). Mu-opioid receptor system modulates responses to vocal bonding and distress signals in humans. Philosophical Transactions of the Royal Society B: Biological Sciences. See also the whole theme issue Cracking the laugh code in Phil Trans B here.
  11. Sun, L., Lukkarinen, L., Noppari, T., Nazari-Farsani, S., Putkinen, V., Seppälä, K., Hudson, M., Tani, P., Lindberg, N., Karlsson, H., Hirvonen, J., Salomaa, M., Venetjoki, N., Lauerma, H., Tiihonen, H., & Nummenmaa, L. (2022). Aberrant motor contagion of emotions in psychopathy and high-functioning autism. Cerebral Cortex.
  12. Laurila, S., Rebelos, E., Lahesmaa, M., Sun, L., Schnabl, K., Peltomaa, T-M., Klen, R., U-Din, M., Honka, M-J., Eskola, O., Kirjavainen, A.K., Nummenmaa, L., Klingenspor, M., Virtanen, K.A., & Nuutila, P. (2022). Novel effects of the gastrointestinal hormone secretin on cardiac metabolism and renal function. American Journal of Physiology-Endocrinology and Metabolism, E54E62, 2022. 
  13. Kantonen, T., Pekkarinen, L., Karjalainen, T., Bucci, M., Kalliokoski, K., Haaparanta-Solin, M., Aarnio, R., Dickens, A.M., von Eyken, A., Laitinen, K., Houttu, N., Kirjavainen, A.K., Helin, S., Hirvonen, J., Nuutila, P., & Nummenmaa, L. (2022). Obesity risk is associated with altered cerebral glucose metabolism and decreased mu-opioid and CB1-receptor availability.  International Journal of Obesity.


  1. Saarimäki, H.P., Glerean, E., Smirnov, D., Jääskeläinen, I.P., Sams, M., & Nummenmaa, L. (2021). Classification of emotions based on functional connectivity patterns of the human brain. NeuroImage.
  2. Nummenmaa, L., Jern, P., Malen, T., Kantonen, T., Pekkarinen, L., Lukkarinen, L., Sun, L., Nuutila, P., & Putkinen, V. (2021). µ-opioid receptor availability is associated with sex drive in human males. Cognitive, Affective, & Behavioral Neuroscience. 
  3. Rantanen M., Hautala, J., Loberg, O., Nuorva, J., Hietanen, J.K., Nummenmaa, L., & Astikainen, P. (2021). Attentional bias to interpersonal aggression in depression – an eye movement study. Scandinavian Journal of Psychology, 62, 639647. 
  4. Kantonen, T. Karjalainen, T., Pekkarinen, L., Isojärvi, J., Kalliokoski, K., Kaasinen, V., Hirvonen, J., Nuutila, P., & Nummenmaa, L. (2021). Cerebral μ-opioid and CB1-receptor systems have distinct roles in human feeding behavior. Translational Psychiatry, 11, 442.
  5. Nummenmaa, L., Lukkarinen, L., Sun, L., Putkinen, V., Seppälä, K., Karjalainen, T., Karlsson, H.K., Hudson, M., Venetjoki, N., Salomaa, M., Rautio, P., Hirvonen, J., Lauerma, H., & Tiihonen, J. (2021). Brain basis of psychopathy in criminal offenders and general population. Cerebral Cortex, 31, 4104–4114.
  6. Laurila, S., Lahesmaa, M., Sun, L., Brain, K., Laitinen, K., Niemi, T., Taittonen, M., Din, M.U., Klen, R., Välikangas, T., Elo, L., Eskola, O., Kirjavainen, A., Nummenmaa, L., Virtanen, K.A., Klingenspor, M., & Nuutila, P (2021). Secretin activates brown fat and induces satiation in humans. Nature Metabolism, 3, 798-809.
  7. Karlsson, H.K., Tuominen, L., Helin, S., Salminen, P., Nuutila, P. & Nummenmaa, L. (2021). Preoperative µ-opioid receptor availability predicts weight loss following bariatric surgery. JCI Insight, 6, e147820
  8. Suvilehto, J.T., Renvall, V., & Nummenmaa, L. (2021). Relationship-specific encoding of social touch in somatosensory and insular cortices. Neuroscience, 364, 105-116.  
  9. Komulainen, E., Glerean, E., Heikkilä, R., Nummenmaa, L., Raij, T., Isometsä, E., & Ekelund, J. (2021). Escitalopram enhances synchrony of brain responses during emotional narratives in patients with major depressive disorder. NeuroImage,118110.  
  10. Karlsson, H.K., Tuominen, L., Helin, S., Salminen, P., Nuutila, P. & Nummenmaa, L. (2021). Mesolimbic opioid-dopamine interaction is disrupted in obesity but recovered by weight loss following bariatric surgery. Translational Psychiatry, 11, 259.
  11. Putkinen, V., Nazari-Farsani, S., Seppälä, K., Karjalainen, T. Sun, L., Karlsson, H., Hudson, M., Heikkilä, T.K., Hirvonen, J., & Nummenmaa, L. (2021). Decoding music-evoked emotions in the auditory and motor cortex. Cerebral Cortex, 31, 2549-2560.
  12. Manninen, S., Karjalainen, T., Tuominen, L.J., Hietala, J., Kaasinen, V., Joutsa, J., Rinne, J., & Nummenmaa, L. (2021). Cerebral gray matter density is associated with neuroreceptor and neurotransporter availability: A combined PET and MRI study. NeuroImage, 235, 117968.
  13. Rebelos, E., Honka, M.J., Ekblad, L., Bucci, M., Hannukainen, J.C., Silva, L.F., Virtanen, K., Nummenmaa, L., & Nuutila, P. (2021). The obesity risk SNP (rs17782313) near the MC4R gene is not associated with brain glucose uptake during insulin clamp in humans. Journal of Clinical Medicine: Clinical Neurology, 10, 1312.
  14. Turtonen, O., Saarinen, A., Nummenmaa, L., Tuominen, L., Tikka, M., Armio, R-L., Hautamäki, A., Laurikainen, H., Raitakari, O., Keltikangas-Järvinen, L., & Hietala, J. (2021). Adult attachment system links with brain μopioid receptor availability in vivo. Biological Psychiatry -Cognitive Neuroscience and Neuroimaging, 6,360-369.
  15. Rebelos, E., Bucci, M., Karjalainen, T., Oikonen, V., Bertoldo, A., Hannukainen, J.C., Virtanen, K.A., Latva-Rasku, A., Hirvonen, J., Heinonen, I., Parkkola, R., Laakso, M., Ferrannini, E., Iozzo, P., Nummenmaa, L., & Nuutila, P. (2021). Insulin resistance is associated with enhanced brain glucose uptake during euglycemic hyperinsulinemia: A large-scale PET cohort. Diabetes Care, 44, 788-794.
  16. Sun, L.. Tang, J., Liljenbäck, H., Honkaniemi, A., Virta, J., Isojärvi, J., Karjalainen, T., Kantonen, T., Nuutila, P., Hietala, J., Kaasinen, V., Kalliokoski, K., Hirvonen, J., Scheinin, H. Helin, S., Eerola, K., Savontaus, E., Yatkin, E., Rinne, J.O., Roivainen, A., & Nummenmaa, L. (2021). Seasonal variation in brain mu-opioid receptor availability. The Journal of Neuroscience, 41, 1265-1273.
  17. Parsons, C., Young, C., Sinerva, E., Kajanoja, J., Nummenmaa, L., Korja R., Karlsson, L., Karlsson, H. (2021).  Investigating the effects of perinatal status and gender on adults’ responses to infant and adult facial emotion. Emotion, 21, 337-349.
  18. Heiskanen, M.A., Honkala, S.M., Ojala, R.O., Lautamäki, R., Koskensalo, K., Lietzén, M., Hentilä, J., Saunavaara, V., Saunavaara, J., Helmiö, M., Löyttyniemi, E., Nummenmaa L., Collado, M.C., Malm, T., Lahti, L., Pietiläinen, K.H., Kaprio, J., Rinne, J.O., & Hannukainen, J.C. (2021). Systemic cross-talk between brain, gut, and peripheral tissues in glucose homeostasis: effects of exercise training (CROSSYS). Exercise training intervention in monozygotic twins discordant for body weight. BMC Sports Science, Medicine and Rehabilitation, 13, 16. 


  1. Volynets, S., Glerean, E., Hietanen, J.K., Hari, R., & Nummenmaa, L. (2020). Bodily maps of emotions are culturally universal. Emotion, 2020. 20, 1127-1136. Dataset may be downloaded here.
  2. Volynets, S., Smirnov, D., Saarimäki, H., & Nummenmaa, L. (2020). Statistical pattern recognition reveals shared neural signatures for displaying and recognizing specific facial expressions. Social Cognitive and Affective Neuroscience.   
  3. Nummenmaa, L., Karjalainen, T., Isojärvi, J., Kantonen, T., Tuisku, J., Kaasinen, V., Joutsa, J., Nuutila, P., Kalliokoski, K., Hirvonen, J., Hietala, J. & Rinne, J. (2020). Lowered endogenous mu-opioid receptor availability in subclinical depression and anxiety. Neuropsychopharmacology, 45, 1953–1959.
  4. Kantonen, T., Karjalainen, T., Isojärvi, J., Nuutila, P., Tuisku, J., Rinne, J., Hietala, J., Kaasinen, V., Kalliokoski, K., Scheinin, H., Hirvonen, J., Vehtari, A., & Nummenmaa, L.  (2020). Interindividual variability and lateralization of mu-opioid receptors in the human brain. NeuroImage, 217, 116922.
  5. Rebelos, E., Hirvonen, J., Bucci, M., Pekkarinen, L., Nyman, M., Hannukainen, J.C., Iozzo, P., Salminen, P., Nummenmaa, L., Ferrannini, E., & Nuutila, P. (2020). Brain FFA uptake is elevated in morbid obesity, and is irreversible six months after bariatric surgery: a positron emission tomography study. Diabetes, Obesity, and Metabolism, 22, 1074–1082. 
  6. Hudson. M., Seppälä, K., Putkinen, V., Sun, L., Glerean, E., Karjalainen, T., Karlsson, H.K., Hirvonen, J., & Nummenmaa, L. (2020). Dissociable neural systems for unconditioned acute and sustained fear. NeuroImage, 216, 116522.
  7. Rebelos, E., Mari, A., Bucci, M., Hannukainen, J.C., Virtanen, K.A., Hirvonen, J., Nummenmaa, L., Heni, M., Iozzo, P., Ferrannini, E., & Nuutila, P. (2020). Brain substrate metabolism and ß-cell function in humans: a positron emission tomography study. Endocrinology, Diabetes & Metabolism, 00:e00136, 1-10.
  8. Renvall, V., Kauramäki, J., Malinen, S., Hari, R., & Nummenmaa, L. (2020). Imaging real-time tactile interaction with two-person dual-coil fMRI. Frontiers in Psychiatry.
  9. Karjalainen, T., Tuisku, J., Santavirta, S., Kantonen, T., Bucci, M., Tuominen, L., Hirvonen, J., Hietala, J., Rinne J., & Nummenmaa, L. (2020). Magia: Robust automated image processing and kinetic modeling toolbox for PET neuroinformatics. Frontiers in Neuroinformatics. You can install Magia to your own system on GitHub
  10. Farsani, S., Nyman, M., Karjalainen, T., Bucci, M., Isojärvi, J., & Nummenmaa, L. (2020). Automated segmentation of acute stroke lesions using a data-driven anomaly detection on diffusion weighted MRI Journal of Neuroscience Methods, 333, 108575.


  1. Karjalainen, T., Seppälä, K., Glerean, E., Karlsson, H.K., Lahnakoski, J.M., Nuutila, P., Jääskeläinen, I-P., Hari, R., Sams, M., Nummenmaa, L. (2019). Opioidergic regulation of emotional arousal: A combined PET–fMRI study. Cerebral Cortex, 14,4006-4016.
  2. Smirnov, D., Saarimäki, H., Glerean, E., Hari, R., Sams, M. & Nummenmaa, L. (2019). Emotions amplify speaker–listener neural alignment. Human Brain Mapping, 40, 4777-4788.
  3. Suvilehto, J.T., Nummenmaa, L., Harada, T., Dunbar, R.I.M., Hari, R., Turner, R., Sadato, N., & Kitada, R. (in press). Cross-cultural similarity in relationship-specific social touching. Proceedings of The Royal Society of London Series B: Biological Sciences, 286, 1-10Data may be downloaded here and code here.
  4. Heiskanen, T., Leppä, M., Suvilehto, J., Akural, E., Larinkoski, T., Jääskeläinen, I.P., Sams, M., Nummenmaa, L., & Kalso, E. (2019). The opioid agonist remifentanil increases subjective pleasure. British Journal of Anaesthesia, 122, 216–219.
  5. Torregrossa, L., Snodgrass, M., Hong, S.-J., Heathman, N., Glerean, E., Nummenmaa, L., & Park, S. (2019) Anomalous bodily maps of emotions in schizophrenia. Schizophrenia Bulletin, 45,1060-1067.
  6. Lähteenmäki, M., Kauramäki, J., Sauter, D., & Nummenmaa, L. (2019). Auditory affective processing requires awareness. Emotion, 19, 53-69.See also our prior study showing similar constraits in visual processing.
  7. Rebelos, E., Immonen, H., Bucci, M., Hannukainen, J.C., Nummenmaa, L., Honka, MJ., Soinio, M., Salminen, P., Ferrannini, E., Iozzo, P., & Nuutila, P. (in press). Endogenous glucose production is associated with brain glucose uptake in morbidly obese subjects. Diabetes, Obesity and Metabolism, 21218-226.
  8. Nyman, M.J., Eskola, O., Kajander, J., Jokinen, R., Penttinen, J., Karjalainen, T., Nummenmaa, L., Hirvonen, J., Burns, D. Hargreaves, R., Solin, O., & Hietala, J. (2019).Brain neurokinin-1 receptor availability in never medicated patients with major depression: A pilot study Journal of Affective Disorders, 242,188-194.


  1. Mäntylä, T., Nummenmaa, L., Rikandi, E., Lindgren, M., Kieseppä, T., Hari, R., Suvisaari, J., & Raij, T. (2018). Aberrant functioning of integrative hubs in first-episode psychosis during natural audiovisual processing. Biological Psychiatry, 84,655-664. See also the commentary for our paper by Gaurav Patel in the same journal.
  2. Nummenmaa, L., Hari, R., Hietanen, J.K., & Glerean, E. (2018). Maps of subjective feelings. Proceedings of the National Academy of Sciences of the United States of America, 115, 9198–9203. Data may be downloaded here and code here.
  3. Saanijoki, T., Nummenmaa, L., Tuulari, J.J.m Tuominen, L., Arponen, E., Kalliokoski, K. & Hirvonen, J. (2018). Aerobic exercise modulates anticipatory reward processing via the μ-opioid receptor system. Human Brain Mapping, 39, 3972-3983.
  4. Gotsopoulos, A., Saarimäki, H., Glerean, E., Jääskeläinen, I.P., Sams, M., Nummenmaa, L., & Lampinen, J. (2018). Reproducibility of importance extraction methods in neural network based fMRI classification. NeuroImage, 181, 44-54. 
  5. Saarimäki, H., Ejthedian, L.F., Glerean, E., Jääskeläinen, I.P., Vuilleumier, P., Sams, M., & Nummenmaa, L. (2018). Distributed affective space represents multiple emotion categories across the brain. Social Cognitive and Affective Neuroscience, 13, 471-482.
  6. Komulainen, E., Heikkilä, R., Nummenmaa, L., Raij, T., Harmer, K., Isometsä, E., & Ekelund, J. (2018). Short-term escitalopram treatment normalizes aberrant self-referential processing in major depressive disorder. Journal of Affective Disorders, 236,  222–229.
  7. Joutsa, J.J., Karlsson, H.K., Majuri, J., Nuutila, P., Helin, S., Kaasinen, V., & Nummenmaa, L. (2018). Binge eating disorder and morbid obesity are associated with lowered mu-opioid receptor availability in the brain. Psychiatry Research: Neuroimaging, 276, 41-45.
  8. Nummenmaa, L., Saanijoki, T., Tuominen, L., Hirvonen, J., Tuulari, J.J., Nuutila, P., & Kalliokoski, K (2018). μ-opioid receptor system mediates reward processing in humans. Nature Communications, 9, 1-7.
  9. Li, J., Oksama, L., Nummenmaa, L. & Hyönä, J. (2018). Angry faces are tracked more easily than neutral faces during multiple identity trackingCognition & Emotion, 32, 464-479.
  10. Saanijoki, T., Nummenmaa, L., Koivumäki, M., Kalliokoski, K.K., Hannukainen, J. (2018). Affective adaptation to repeated SIT and MICT protocols in insulin resistant subjectsMedicine & Science in Sports & Exercise, 50,18–27.
  11. Latva-Rasku, A., Honka, M-J., Stancakova, A., Koistinen, H., Kuusisto, J., Guan, L., Manning, A., Stringham H., Gloyn, A., Lidgren, C., Collins, F., Mohlke, K, Scott, L., Karjalainen, T., Nummenmaa, L., Nuutila, P., & Laakso, M. (2018). A partial loss-of-function variant in AKT2 is associated with reduced insulin-mediated glucose uptake in multiple insulin sensitive tissues: a genotype-based callback positron emission tomography study. Diabetes, 67, 334-342.
  12. Saanijoki, T., Tuominen, L.J., Nummenmaa, L., Arponen, E., Kalliokoski, K., & Hirvonen, J. (2018). Opioid release after high-intensity interval training in healthy human subjectsNeuropsychopharmacology, 43, 246-254.


  1. Smirnov, D., Lachat, F., Peltola, T., Lahnakoski, J.M., Koistinen, O-P., Gleran, E., Vehtari, A., Hari, R., Sams, L., & Nummenmaa, L. (2017). Brain-to-brain hyperclassification reveals action-specific motor mapping of observed actions in humans. PLoS One.
  2. Tuulari, J.J., Tuominen, L., de Boer, F., Hirvonen, J., Nuutila, P., & Nummenmaa, L. (2017). Feeding releases endogenous opioids in humans. The Journal of Neuroscience, 37, 8284-8291.
  3. Karjalainen, T., Karlsson, H.K., Lahnakoski, J.M., Glerean, E., Nuutila, P., Jääskeläinen, I.P., Hari, R., Sams, M., & Nummenmaa, L. (2017). Dissociable roles of cerebral μ-opioid and type 2 dopamine receptors in vicarious pain: A combined PET–fMRI study. Cerebral Cortex, 27, 4257–4266.
  4. Lahnakoski, J., Jääskeläinen, I.P., Sams, M., & Nummenmaa, L. (2017). Neural mechanisms for integrating consecutive and interleaved natural events. Human Brain Mapping, 38, 3360-3376.
  5. Salmi, J., Koistinen, O-P., Jylänki, P., Vehtari, A., Jääskeläinen, I-P., Mäkelä, S., Nummenmaa, L., Nummi-Kuisma, K., Nummi, I., & Sams, M. (2017). Distributed neural signatures of natural audiovisual speech and music in the human auditory cortex. NeuroImage, 157, 108-117.
  6. Manninen, S., Tuominen, L., Dunbar, R.I.M., Karjalainen, T., Hirvonen, J., Arponen, E., Hari, R., Jääskeläinen, I.P., Sams, M., & Nummenmaa, L. (2017). Social laughter triggers endogenous opioid release in humans. The Journal of Neuroscience, 37, 6125-6131; see also commentary on our work by Fausto Caruana in the same journal
  7. Komulainen, E., Glerean, E., Meskanen, K., Heikkilä, R., Nummenmaa, L., Raij, T.T., Lahti, J., Jylhä, P., Melartin, T., Isometsä, E., & Ekelund, J., (2017). Single dose of mirtazapine modulates whole-brain functional connectivity during emotional narrative processing. Psychiatry Research: Neuroimaging, 30, 60-69.
  8. Nummenmaa, L., Oksama, L., Glerean, E., & Hyönä, J. (2017). Cortical circuit for binding object identity and location during multiple object tracking. Cerebral Cortex, 27, 162–172.
  9. Chen,T.,  Nummenmaa, L.,  & Hietanen, J.K. (2017). Eye contact judgment is influenced by perceiver’s social anxiety but not by their affective stateFrontiers in Psychology.


  1. Hautala, J. Lohberg, O., Hietanen, J.K., Nummenmaa, L., & Astikainen, P. (2016). Effects of conversation content on viewing dyadic conversations. Journal of Eye Movement Research, 9, 1-12.
  2. Nummenmaa, L., Tuominen, L., Dunbar, R., Hirvonen, J., Manninen, S., Arponen, E., Machin, A., Hari, R., Jääskeläinen, I.P., & Sams, M. (2016). Social touch modulates endogenous µ-opioid system activity in humansNeuroImage, 138, 242-247.
  3. Tuulari, J.J., Karlsson, H.K., Antikainen, O., Hirvonen, J., Pham, T., Salminen, P., Helmiö, M., Parkkola, R., Nuutila, P., & Nummenmaa, L. (2016). Bariatric surgery induces white and grey matter density recovery in the morbidly obese: A voxel-based morphometric studyHuman Brain Mapping, 37, 3745-3756.
  4. Karjalainen, T., Tuominen, L., Manninen, S., Kalliokoski, K., Nuutila, P., Jääskeläinen, I.P., Hari, R., Sams, M., & Nummenmaa, L. (2016). Behavioural activation system sensitivity is associated with cerebral μ-opioid receptor availabilitySocial Cognitive and Affective Neuroscience, 11, 1310-1316.
  5. Hietanen, J.K., Glerean, E., Hari, R., & Nummenmaa, L. (in press). Bodily maps of emotions across child developmentDevelopmental Science.
  6. Karlsson, H.K., Tuulari, J.J., Tuominen, L., Hirvonen, J., Honka, H., Parkkola, R., Helin, S., Salminen, P., Nuutila, P., & Nummenmaa, L. (2016). Weight loss after bariatric surgery normalizes brain opioid receptors in morbid obesity. Molecular Psychiatry, 21, 1057-1062.
  7. Saarimäki, H., Gostopoulos, A., Jääskeläinen, I.P., Lampinen, J., Vuilleumier, P., Hari, R., Sams, M., & Nummenmaa, L. (2016). Discrete neural signatures of basic emotionsCerebral Cortex, 6, 2563-2573.
  8. Nummenmaa, L., Suvilehto, J.T., Glerean, E., Santtila, P., & Hietanen, J.K. (2016). Topography of human erogenous zonesArchives of Sexual Behaviour, 45, 1207-1216.
  9. Glerean, E., Pan, R., Salmi, J., Kujala, R., Roine, U., Nummenmaa, L., Leppämäki, S., Nieminen-von Wendt, Tani, P., Saramäki, J., Sams, M., & Jääskeläinen, I.P. (2016). Reorganization of functionally connected brain subnetworks in high-functioning autismHuman Brain Mapping, 37, 1066–1079.
  10. Komulainen, E., Heikkilä, R., Meskanen, K., Raij, T.T, Nummenmaa, L., Lahti, J., Jylhä, P, Melartin, T., Isometsä, E., & Ekelund, J. (2016) A single dose of mirtazapine attenuates the default mode network activity during self-referential processing in healthy volunteers. Journal of Psychopharmacology30, 23-32.


  1. Saanijoki, T., Nummenmaa, L., Eskelinen, J.J., Savolainen, A.M., Vahlberg, T., Kalliokoski, K.K., & Hannukainen, J.C. (2015). Affective responses to repeated sessions of high-intensity interval trainingMedicine & Science in Sports & Exercise,47, 2604-2611.
  2. Suvilehto, J., Glerean, E., Dunbar, R.I.M., Hari, R., & Nummenmaa, L.
    (2015).Topography of social touching depends on emotional bonds between humans.Proceedings of the National Academy of Sciences of the United States of America, 115, 13811-13816.
  3. Tuominen, L., Tuulari, J.J., Karlsson, H.K.,  Hirvonen, J., Helin, S., Salminen, P., Parkkola, R., Hietala, J., Nuutila, P., & Nummenmaa, L. (2015). Aberrant mesolimbic dopamine-opiate interaction in obesity. NeuroImage122, 80-86.
  4. Hotta, J., Harno, H., Nummenmaa, L., Hari, R., & Forss, N. (2015). Patients with complex regional pain syndrome overestimate applied force in observed hand actions. European Journal of Pain, 19, 1372-1381.
  5. Nummenmaa, L., Manninen, S., Tuominen, L., Hirvonen, J., Kalliokoski, K., Nuutila, P., Jääskeläinen, I.P. Hari, R., Dunbar, R.I.M., & Sams, M. (2015). Adult attachment style is associated with cerebral mu-opioid receptor availability in humansHuman Brain Mapping, 36, 3621-3628.
  6. Halko, M-L., Mäkelä, T., Nummenmaa, L., Hlushchuk, Y., & Schürmann, M. (2015).Hedonic context modulates risky choices and reward responses in amygdala and dorsal striatum. Journal of Neuroscience, Psychology and Economics8, 100-115.
  7. Alho, J., Salminen, N., Sams, M., Hietanen, J.K. & Nummenmaa, L. (2015).Facilitated early cortical processing of nude human bodiesBiological Psychology,109,103–110.
  8. Lähteenmäki, M., Hyönä, J., Koivisto, M., & Nummenmaa, L. (2015). Affective processing requires awarenessJournal of Experimental Psychology: General144, 339-365.
  9. Karlsson, H.K., Tuominen, L., Tuulari, J.J., Hirvonen, J., Parkkola, R., Helin, S., Salminen, P., Nuutila, P., & Nummenmaa, L. (2015). Obesity is associated with decreased mu-opioid but unaltered dopamine D2 receptor availability in the brain.The Journal of Neuroscience35, 3959-3965.
  10. Tuulari, J.J., Karlsson, H.K., Hirvonen, J., Salminen, P., Nuutila, P., & Nummenmaa, L. (2015). Neural circuits for cognitive appetite control in healthy and obese individuals: An fMRI studyPLoS One10(2), e0116640.


  1. Calvo, M.G., Gutiérrez-García, A., Fernández-Martín, A., & Nummenmaa, L. (2014).Recognition of facial expressions of emotion is related to their frequency in everyday lifeJournal of Nonverbal Behavior38, 549-567.
  2. Smirnov, D., Glerean, E., Lahnakoski, J., Salmi, J., Jääskeläinen, I.P., Sams, M.,Nummenmaa, L. (2014). Fronto-parietal network supporting context-dependent speech comprehensionNeuropsychologia, 63, 293–303.
  3. Hietanen, J.K., Kirjavainen, I., & Nummenmaa, L., (2014). Additive effects of affective arousal and top-down attention on the event-related brain responses to human bodiesBiological Psychology, 103, 167–175.
  4. Nummenmaa, L., Saarimäki, H., Glerean, E., Gotsopoulos, A., Hari, R., & Sams, M. (2014). Emotional speech synchronizes brains across listeners and engages large-scale dynamic brain networks. NeuroImage, 102, 498-509.
  5. Lahnakoski, J., Glerean, E.G., Jääskeläinen, I.P., Hyönä, J., Hari, R., Sams, M., &Nummenmaa, L. (2014). Synchronous brain activity across individuals underlies shared psychological perspectivesNeuroImage, 100, 316–324.
  6. Orava, J., Nummenmaa, L.,Noponen, T., Viljanen, T., Parkkola, R., Nuutila, P. & Virtanen, K. (2014). Brown adipose tissue function is accompanied by cerebral activation in lean humansJournal of Cerebral Blood Flow and Metabolism, 34, 1018-23.
  7. Tuominen, L., Nummenmaa, L., Keltikangas-Järvinen, L., Raitakari, O. & Hietala, J. (2014). Mapping neurotransmitter networks with PET: An example on serotonin and opioid systemsHuman Brain Mapping, 35, 1875-1884.
  8. Calvo, M.G., Fernández-Martín, A., & Nummenmaa, L. Facial expression recognition in peripheral versus central vision:  Role of the eyes and the mouth (2014).Psychological Research, 78, 180-195.
  9. Nummenmaa, L., Smirnov, D., Lahnakoski, J., Glerean, E., Jääskeläinen, I.P., Sams, M., & Hari, R. (2014). Mental action simulation synchronizes action-observation circuits across individualsThe Journal of Neuroscience, 34,748 –757.
  10. Nummenmaa, L., Glerean, E., Hari, R., & Hietanen, J.K. (2014). Bodily maps of emotionsProceedings of the National Academy of Sciences of the United States of America, 111, 646-651.


  1. Karlsson, H.K., Tuulari, J.J., Hirvonen, J., Lepomäki, V., Parkkola, R., Hiltunen, J., Hannukainen, J., Nuutila, P., & Nummenmaa, L. (2013). Obesity is associated with white matter atrophy: A combined diffusion tensor imaging and voxel-based morphometric studyObesity, 21, 2530-2537.
  2. Salmi, J., Roine, U., Glerean, E., Lahnakoski, J., Nieminen-von Wendt, T., Tani, P., Leppämäki, S., Nummenmaa, L., Jääskeläinen, I.P., Carlson, S., Rintahaka, P., & Sams, M. (2013). The brains of high functioning autistic individuals do not synchronize with those of othersNeuroImage: Clinical3, 489–497.
  3. Kätsyri, J., Ravaja, N., Hari, R., & Nummenmaa, L. (2013). The opponent matters: Elevated fMRI reward responses to winning against a human versus a computer opponent during interactive video game playingCerebral Cortex23,2829-2839.
  4. Calvo, M.G., Fernández-Martín, A., & Nummenmaa, L. (2013) A smile biases the recognition of eye expressions: Configural projection from a salient mouthQuarterly Journal of Experimental Psychology. 66, 1159-1181.
  5. Tuulari, J.J., Karlsson, H.K., Hirvonen, J., Bucci, M., Hannukainen, J.M., salminen, P., Helmiö, M., Ovaska, J., Soinio, M., Forsback, S., Nummenmaa, L., & Nuutila, P.Weight loss after bariatric surgery reverses insulin-induced increases in brain glucose metabolism of the morbidly obese (2013). Diabetes, 62, 2747-2751.
  6. Kätsyri, J., Ravaja, N., Hari, R., & Nummenmaa, L. (2013). Just watching the game ain’t enough: Striatal fMRI reward responses to successes and failures in a video game during active and vicarious playingFrontiers in Human Neuroscience.


  1. Calvo, M.G., Fernández-Martín, A., & Nummenmaa, L. (2012).  Perceptual and affective processing of ambiguous smiling facial expressions. Cognition, 125, 373-393.
  2. Laaksonen, K., Kirveskari, E. Mäkelä, J.P., Kaste, M., Mustanoja, S., Tatlisumak, T,Nummenmaa, L., & Forss, N. Effect of afferent input on motor cortex excitability during stroke recovery (2012). Clinical Neuropshysiology123, 2429-2436.
  3. Nummenmaa, L., Hietanen, J.K., Santtila, P., & Hyönä, J. (2012). Gender and visibility of sexual cues modulate eye movements while viewing faces and bodies.Archives of Sexual Behavior, 41, 1439-1451.
  4. Viinikainen, M., Glerean, E., Jääskeläinen, I.P., Kettunen, J., Sams, M., &Nummenmaa, L. (2012). Nonlinear neural representation of emotional feelings elicited by dynamic naturalistic stimulationThe Open Neuroimaging Journal, 2.
  5. Lahnakoski, J.M., Glerean, E., Salmi, J., Jääskeläinen, I.P., Sams, M., Hari, R. &Nummenmaa, L. (2012). Naturalistic fMRI mapping reveals superior temporal sulcus as the hub for the distributed brain network for social perceptionFrontiers in Human Neuroscience.
  6. Nummenmaa, L., Glerean, E., Viinikainen, M., Jääskeläinen, I.P., Hari, R., & Sams, M. (2012) Emotions promote social interaction by synchronizing brain activity across individuals. Proceedings of the National Academy of Sciences of the United States of America, 109, 9599–9604.
  7. Nevalainen, P., Pihko, E., Mäenpää, H., Valanne, L., Nummenmaa, L., & Lauronen, L. (2012). Bilateral alterations in somatosensory cortical processing in hemiplegic cerebral palsy. Developmental Medicine and Child Neurology, 54, 361-7.
  8. Nummenmaa, L., Engell, A.D., von dem Hagen, E., Henson, R.N.A., & Calder, A.J. (2012). Autism  spectrum  traits  predict  the  neural  response  to  eye  gaze  in  typical  individuals. NeuroImage, 59, 3356-3363.
  9. Nummenmaa, L., Hirvonen, J., Hannukainen, J.C, Immonen, H., Lindroos, M.M., Salminen, P., & Nuutila, P. (2012). Dorsal striatum and its limbic connectivity mediate abnormal anticipatory reward processing in obesityPLoS One, 7(2), e31089.


  1. Calvo, M.G. & Nummenmaa, L. (2011). Time course of discrimination between emotional facial expressions: The role of visual saliencyVision Research, 51, 1751–1759.
  2. Calvo, M.G., Avero, P., & Nummenmaa, L. (2011). Primacy of emotional vs. semantic scene recognition in peripheral visionCognition & Emotion, 25,1358-75.
  3. Hietanen, J.K. & Nummenmaa, L. (2011). The Naked Truth: The Effect of Nudity on Face and Body Sensitive N170 ResponsePLoS One, 6(11), e24408.
  4. Hirvonen J., Virtanen, K.A., Nummenmaa, L., Hannukainen, J.C., Honka, M-J., Bucci, M., Nesterov, S.V., Parkkola, R., Rinne, J., Iozzo, P., & Nuutila, P. Effects of insulin on brain glucose metabolism in impaired glucose tolerance. (2011).Diabetes60, 443-447.
  5. Kallio, S., Hyönä, J., Revonsuo, A., Pilleriin, S., & Nummenmaa, L. (2011). The existence of a hypnotic state revealed by eye movements. PLoS One6, e26374 – e26374.
  6. Nummenmaa, L., Hietanen, J.K., Calvo, M.G. & Hyönä, J. (2011). Food catches the eye but not for everyone: A BMI–contingent attentional bias in rapid detection of nutrimentsPLoS One, 6(5), e19215.
  7. von dem Hagen, E., Nummenmaa, L., Yu, R., Engell, A., Ewbank, M., &  Calder, A. (2011). Autism Spectrum traits in the typical population predict structure and function in the posterior superior temporal sulcusCerebral Cortex, 21, 493-500


  1. Calvo, M.G., Nummenmaa, L., & Avero, P. (2010). Recognition advantage of happy faces in extrafoveal vision: Featural and affective processingVisual Cognition, 18, 1274 – 1297.
  2. Engell, A.D., Nummenmaa, L., Oosterhof, N.N., Henson, R.N.A., Haxby, J.V., & Calder, A.J. (2010). Differential activation of frontoparietal attention networks by social and symbolic spatial cuesSocial Cognitive and Affective Neuroscience, 5,432-440.
  3. Nummenmaa, L., Passamonti, L., Rowe, J., Engell, A.D., & Calder, A.J. (2010)Connectivity analysis reveals a cortical network for eye gaze perception. Cerebral Cortex8, 1780-1787.
  4. Nummenmaa, L., & Hyönä, J., & Calvo, M.G. (2010). Semantic categorization precedes affective evaluation of visual scenesJournal of Experimental Psychology: General, 139, 222-246.


  1. Calvo, M.G. & Nummenmaa, L. (2009). Eye movement assessment of the time course in facial expression recognition: Neurophysiological implications.Cognitive, Affective and Behavioral Neuroscience, 9, 398-411.
  2. Calvo, M.G. & Nummenmaa, L. (2009). Lateralised covert attention in word identification. Laterality, 14(2),178- 195.
  3. Gutierrez, A., Nummenmaa, L., & Calvo, M.G. (2009). Enhanced processing of a coarse impression of emotional scenes in peripheral visionSpanish Journal of Psychology, 12, 414-423.
  4. Nummenmaa, L. & Hietanen, J. K (2009). How attentional systems process conflicting cues? The superiority of social over symbolic orienting revisited.Journal of Experimental Psychology: Human Perception and Performance35, 1738-1754.
  5. Nummenmaa, L., Hyönä, J., & Calvo, M.G. (2009). Emotional scene content drives the saccade generation system reflexively. Journal of Experimental Psychology: Human Perception and Performance, 35, 305-323.
  6. Nummenmaa, L., Hyönä, J., & Hietanen, J.K. (2009). I’ll walk this way: Eyes reveal the direction of locomotion and make passers-by to look and go the other wayPsychological Science20, 1454-1458.


  1. Calvo, M.G. & Nummenmaa, L. (2008). Detection of emotional faces: salient physical features guide effective visual search. Journal of Experimental Psychology: General, 137, 471-494.
  2. Calvo, M.G., Nummenmaa, L., & Avero, P. (2008). Visual search of emotional faces: Eye-movement assessment of component processesExperimental Psychology55, 359-370.
  3. Calvo, M.G., Nummenmaa, L., & Hyönä, J. (2008). Emotional scenes in peripheral vision: Selective orienting and gist processing, but not content identificationEmotion8, 68–80.
  4. Hietanen, J.K., Leppänen, J.M., Nummenmaa, L., & Astikainen, P. (2008).Visuospatial attention shifts by gaze and arrow cues: An ERP study. Brain Research, 1215, 123 – 136.
  5. Nummenmaa, L., Hirvonen, J., Parkkola, R., & Hietanen, J. K. (2008). Is emotional contagion special? An fMRI study on neural systems for affective and cognitive empathyNeuroImage, 43, 571–580.
  6. Nummenmaa, M., & Nummenmaa, L. (2008). Emotions and interest in visible and invisible web-based learning activitiesBritish Journal of Educational Psychology, 78, 163 – 178.
  7. Nummenmaa, L., Peets, K., & Salmivalli, C. (2008). Automatic activation of adolescents’ peer-relational schemas: Evidence from priming with facial identity.Child Development, 79, 1659 – 1675.


  1. Calvo, M.G., Nummenmaa, L., & Hyönä, J. (2007). Emotional and neutral scenes in competition: orienting, efficiency, and identificationThe Quarterly Journal of Experimental Psychology60, 1585-1593.
  2. Calvo, M.G., & Nummenmaa, L. (2007). Processing of unattended emotional visual scenes. Journal of Experimental Psychology: General, 136, 347–369.


  1. Hietanen, J.K., Nummenmaa, L., Nyman, M., Parkkola, R., & Hämäläinen, H. (2006) Automatic attention orienting by social and symbolic cues activate different neural networks: An fMRI studyNeuroImage, 33, 406-413.
  2. Nummenmaa, L., & Hietanen, J.K. (2006) Gaze distractors influence saccadic curvature: Evidence for the role of the oculomotor system in gaze-cued orienting.Vision Research46, 3674–3680.
  3. Nummenmaa, L., & Hyönä, J., & Calvo, M.G. (2006). Eye movement assessment of selective attentional capture by emotional picturesEmotion, 6, 257-268.


  1. Vuorela, M., & Nummenmaa, L. (2004). How undergraduate students meet a new learning environment? Computers in Human Behavior, 20, 763-777.
  2. Vuorela, M., & Nummenmaa, L. (2004). Affect, regulation and behavior in web-based learning environmentsEuropean Journal of Psychology of Education, 19, 423-436.

Review articles 

  1. Knuuti, J., Tuisku, J., Kärpijoki, H., Iida, H., Maaniitty, T., Latva-Rasku, A., Oikonen, V., Nesterov, S., Teuho, J., Jaakkola, M., Klen, R., Louhi, H., Saunavaara, V., Nuutila, P., Saraste, A., Rinne, J., & Nummenmaa, L. (2023). Quantitative perfusion imaging with total body positron emission tomography. Journal of Nuclear Medicine, 64, 11-19.
  2. Pak, K., & Nummenmaa, L. (2023). Brain dopamine receptor system is not altered in obesity: Bayesian and frequentist meta-analyses. Human Brain Mapping, 44, 6552-6560.
  3. Nummenmaa, L. & Van Dillen, L. (2021). Carnal pleasures. Current Opinion in Behavioural Sciences, 39, 85-92.
  4. Nummenmaa, L., Putkinen, V., & Sams, M. (2021). Social pleasures of music. Current Opinion in Behavioral Sciences, 39, 196-202.
  5. Sander, D., & Nummenmaa, L. (2021). Reward and emotion: An affective neuroscience approach. Current Opinion in Behavioural Sciences39, 161–167.
  6. Rebelos, E., Nummenmaa, L., Dadson, P., Latva-Rasku, A., & Nuutila, P. (2020). Brain insulin sensitivity is linked to body fat distribution– the positron emission tomography perspective. European Journal of Nuclear Medicine and Molecular Imaging.
  7. Nummenmaa, L., & Saarimäki, H. (2019). Emotions as discrete patterns of systemic activity. Neuroscience Letters, 6, 3-8.
  8. Nummenmaa, L., & Tuominen, L.J. (2018). Opioid system and human emotions. British Journal of Pharmacology, 175, 2735-2749.
  9. Nummenmaa, L., Lahnakoski, J., & Glerean, E. (2018). Sharing the social world via intersubject neural synchronization. Current Opinion in Psychology, 24, 7-14.
  10. Nummenmaa, L., & Karjalainen, T. (2017). Opioidergic regulation of pain and pleasure in human social relationships. Neuropsychopharmacology, 43, 217–218.
  11. Hari, R, Sams, M., & Nummenmaa, L. (2016). Attending and neglecting people: Bridging neuroscience, psychology and sociologyPhilosophical Transactions of The Royal Society of London Series B: Biological Sciences, 371.
  12. Adolphs, R., Nummenmaa, L., Todorov, A., & Haxby, J.V. (2016). Data-driven approaches in the investigation of social perceptionPhilosphical Transactions of The Royal Society of London Series B, 371
  13. Calvo, M.G. & Nummenmaa, L. (2016). Perceptual and affective mechanisms in facial expression recognition: An integrative review. Cognition & Emotion, 30, 1081-1106.
  14. Nummenmaa, L., & Calvo, M.G. (2015). Dissociation between recognition and detection advantage for facial expressions: A meta-analysisEmotion,15, 243-256.
  15. Nummenmaa, L., & Calder, A.J. (2009). Neural mechanisms of social attention.Trends in Cognitive Sciences, 13(3), 135-143.
  16. Nummenmaa, L., & Niemi, P. (2004). Inducing affective states with success-failure manipulations: A meta-analysis. Emotion, 4, 207-214.

Review articles in Finnish

  1. Nummenmaa, L. & Nuutila, P. (2019). Lihavuusleikkaus parantaa aivojen terveyttä. Duodecim.
  2. Nummenmaa, L. (2016). Aivot, mielihyvä ja liikuntaLiikunta & Tiede, 63, 39-43.
  3. Nummenmaa, L. (2016). Tunteiden neurobiologiaLääkärilehti, 10/2016.
  4. Nummenmaa, L. (2011). Sinun silmiesi tähden: Katseen havaitsemisen aivomekanismit. Psykologia, 46, 4-19.
  5. Nummenmaa, L. (2005). Efektikoko psykologisessa tutkimuksessa [Effect size in psychological science]. Psykologia, 5-6/05, 559-567.
  6. Juujärvi, P., & Nummenmaa, L. (2003). Emootiot, emootion säätely ja hyvinvointi[Emotions, emotion regulation and well being]. Psykologia, 1/2004, 59-66.

Book chapters and non-reviewed articles

  1. Nummenmaa, L., & Hirvonen, J. (in press). Mapping the human emotion circuits with positron emission tomography. in Armony, J., & Vuilleumier, P. The Cambridge Handbook of Human Affective Neuroscience. Cambridge: Cambridge University Press.
  2. Nummenmaa, L., Seppälä. L., & Putkinen, V. (2023). Molecular imaging of the human emotion circuit. In Boggio, P., Wingenbach, T.S., Comfort, W., Lira. M., & Marrins, L.  (Eds.).  Social and Affective Neuroscience of Everyday Human Interaction – From Theory to Methodology. Springer Nature: Cham, Switzerland. You can also download the full ebook here
  3. Nummenmaa, L., & Saarimäki, H. (2021). Tunteelliset aivot. In R.Hari & S. Carlson (Eds.). Aivoaakkoset. Helsinki: Aalto ARTS Books.
  4. Nummenmaa, L, (2020). Psychology and neurobiology of horror movies. Samsung Nordic, Stockholm, Sweden.
  5. Nummenmaa, L., Tuominen, L.J., & Hirvonen, J. (2018). Simultaneous PET-MRI confirms that cerebral blood flow does not confound PET neuroreceptor activation studies. ACS Chemical Neuroscience, 21, 159-161.
  6. Nummenmaa, L. (2017). Mistä puhumme kun puhumme tunteista. Tieteessä tapahtuu, 2/2017, 35-39.
  7. Nummenmaa, L. (2016). Keholliset tunteet. Psykologianopettaja 1/2016, 7-10.
  8. Nummenmaa, L. & Hari, R. (2017). Koskettavat tunteet. In Ylikoski, M. (eds), Tutkitusti-Ihmisen Mieli.
  9. Hari, R., Himberg, T., Nummenmaa, L., Hämäläinen, M., & Parkkonen, L. (2013).Synchrony of brains and bodies during implicit interpersonal interactionTrends in Cognitive Sciences, 17, 105-106.
  10. Hari, R., & Nummenmaa, LMiten toivo ja epätoivo tarttuvat? In M. O. Huttunen, R. Pelkonen, & K. Saarelma (2014). Toivo lääketieteessä. Helsinki: Duodecim.
  11.   Nummenmaa, L. (2013). Tunteet kriisien keskellä. In J. Hämeen-Anttila, K. Katajala, A. Sihvola, & I. Hetemäki. (eds.) Kaikki syntyy kriisistä. Helsinki: Gaudeamus.
  12. Nummenmaa L. & Sams, M. (2011). Tunteet mielessä ja aivoissa. In L.C. Andersson, I. Hetemäki, R. Mustonen & A.Sihvola (eds.). Kaikki irti arjesta. Helsinki: Gaudeamus
  13. Sams, M. & Nummenmaa, L. (2011). Miten suomalaiset kuvailevat tunteitaan? In L.C. Andersson, I. Hetemäki, R. Mustonen & A.Sihvola (eds.). Kaikki irti arjesta. Helsinki: Gaudeamus.
  14. Nummenmaa L. (2009). Tunteiden aivomekanismit. In M. Sams (Eds.). Aivot ja oppiminen. Electronic textbook available at
  15. Nummenmaa, L., Hyona, J., & Calvo, M.G. (2008). Do emotional scenes catch the eye? In K. Rayner, J. Shen, X. Bai, and G. Yan (Eds.), Cognitive and Cultural Influences on Eye Movements.  Tinajin: Tianjin People’s Publishing House/Hove: Psychology Press.
  16. Calder, A.J., & Nummenmaa, L. (2007). Face cells: Separate processing of expression and gaze in the amygdalaCurrent Biology17, R371-R372.
  17. Nummenmaa, L. (2007). Sosiaalisen tarkkaavaisuuden suuntautuminen.Psykologia, 42, 289-294.
  18. Nummenmaa, L. (2006). Emootiotutkimuksen menetelmät [Research methods in Emotion]. In H. Hämäläinen, A. Revonsuo, O. Aaltonen, & M. Laine (Eds.), Mieli ja aivot. Kognitiivisen neurotieteen oppikirja. Jyväskylä: Gummerus.
  19. Nummenmaa, L. (2006) Kogntio ja emootio [Cognition and Emotion]. In H. Hämäläinen, A. Revonsuo, O. Aaltonen, & M. Laine, Mieli ja aivot. Kognitiivisen neurotieteen oppikirja. Jyväskylä: Gummerus. 


  1. Nummenmaa, L. (2023) Käyttäytymistieteiden tilastolliset menetelmät – digikirja. [Multivariate statistics for behavioural sciences – Digital edition]. Helsinki: Tammi.
  2. Heikkilä, T., Nummenmaa, L., Nummenmaa, M., & Tavast, M. (2021). Tunteet ja mielenterveys [Emotions, psychological well being and mental health]. Helsinki: Studeo.
  3. Nummenmaa, L. (2021). Tilastotieteen käsikirja [Handbook of Statistics]. Helsinki: Tammi.
  4. Heikkilä, T., Nummenmaa, L., Nummenmaa, M., Salenius, H. & Tavast, M. (2018). Känslor, psykiskt välbefinnande och mental hälsa [Emotions, psychological well being and mental health]. Helsinki: Tabletkoulu.
  5. Heikkilä, T., Nummenmaa, L., Nummenmaa, M., & Tavast, M. (2017). Tunteet, psyykkinen hyvinvointi ja mielenterveys [Emotions, psychological well being and mental health]. Helsinki: Tabletkoulu.
  6. Nummenmaa, L,, Holopainen, M., & Pulkkinen, P. (2014). Tilastollisten menetelmien perusteet – opettajan opas [Teacher’s guide to Introductory Statistics]. Helsinki: SanomaPRO.
  7. Nummenmaa, L,, Holopainen, M., & Pulkkinen, P. (2014). Tilastollisten menetelmien perusteet [Introductory Statistics]. Helsinki: SanomaPRO.
  8. Nummenmaa, L. (2006; 2009). Käyttäytymistieteiden tilastolliset menetelmät: Harjoituskirja [Excercises on multivariate statistics for behavioural sciences].Vammala: Tammi. Kirja on ladattavissa ilmaiseksi tästä linkistä.
  9. Nummenmaa, L. (2004;2009;2023) Käyttäytymistieteiden tilastolliset menetelmät[Multivariate statistics for behavioural sciences]. Vammala: Tammi.
  10. Nummenmaa, L. (2002). Psykometriikan perusteet [Introduction to psychometrics]. Turku: Turun yliopisto, Psykologian laitos.

Study materials

  1. Vuorela, M., & Nummenmaa, L. (2003) Tiedon hakeminen verkosta. [Information retrieval with digital platforms]. University of Turku: Department of Education.

Books popularizing science

  1. Nummenmaa, L. (2022). Pahuus – ihmisluonnon pimeä puoli (Evil: The Dark Side of Human Nature). Helsinki: Tammi.
  2. Nummenmaa, L. (2018). Tunnekartasto (The Emotion Atlas). Helsinki: Tammi.
  3. Nummenmaa, L. (2010). Tunteiden psykologia (The Psychology of Emotions). Helsinki: Tammi.

Articles popularizing science

  1. Nummenmaa, L. & Hietanen, J.K. (2023). Vitutus. Tehohoitolehti, 2/2023.
  2. Nummenmaa, L. (2023). Tilasto näkee tulevaan. Tiede, 8/2023. (Reprinted on Nov 11th 2023 in Helsingin Sanomat).
  3. Nummenmaa, L. (2022). Vailla omaatuntoa. Tiede, 4, 2022.
  4. Nummenmaa, L. (2020). Mistä tietoisuus syntyy? Tiede, 6, 2020.
  5. Nummenmaa, L. (2019). Häpeä. Suomen Kuvalehti, 16-17/2019, 45.
  6. Hietanen, J.K. & Nummenmaa, L. (2016). Lasten keholliset tunnekokemukset. Psykologi, 06/2016.
  7. Nummenmaa, L.(2016). Keholliset tunteet. Psykologianopettaja 1/2016, 7
  8. Nummenmaa, L. (2016). Tunteet ovat osa jokapäiväistä elämäämme. Potilaan lääkärilehti, 17.3.2016.
  9. Nummenmaa, L. (2015). Tunteet sosiaalisen ihmisen palveluksessaMielenterveys, 2 / 2015.
  10. Nummenmaa, L. (2015). Tunteet, järki ja päätöksentekoChydenius 3 / 2015.
  11. Nummenmaa, L., & Hari, R (2014). Aivot rakastavat seuraaTiede 4 / 2014.
  12. Nummenmaa, L., & Hyönä, J. (2005). Mitä sinä näet tässä kuvassa? Projektiivisen testaamisen tieteellinen perusta [Tell me what do you see in this picture. What is the scientific evidence for validity of projective testing?].Psykologi, 4 / 2005.

PhD theses completed in the Emotion lab

  1. Pekkarinen, L. (2023). Insulin sensitivity and endocannabinoid function as risk factors for obesity: Positron emission tomography studies of insulin sensitivity and CB1 receptorsAnnales Universitatis Turkuensis D 1767.
  2. Kantonen, T. (2022). Brain opioid and endocannabinoid systems as risk factors for obesity. Positron emission tomography studies of µ-opioid and CB1 receptors with glucose uptake analysisAnnales Universitatis Turkuensis D 1625.
  3. Farsani, S.N. (2020). Radiostereometric analysis of initial femoral stem migrations in cementless total hip arthroplasty of postmenopausal women: Exploring contributing factors in initial femoral stem migration. Annales Universitatis Turkuensis D 1511.
  4. Manninen, S. (2019). Endogenous opioid system and human sociability. Annales Universitatis Turkuensis D 1442.
  5. Karjalainen, T. (2019). Opioidergic regulation of human affiliative behavior – Evidence from positron emission tomography studiesAalto University Doctoral Dissertations, 52/2019
  6. Suvilehto, J.T. (2018). Maintaining social bonds via touching: A cross-cultural studyAalto University Doctoral Dissertations, 210/2018.
  7. Saanijoki, T. (2018). Neurobiology of physical exercise: perspectives on psychophysiological effects and opioidergic neurotransmissionAnnales Universitatis Turkuensis D 1363.
  8. Smirnov, D. (2018). Brain basis of sharing and transmitting representations of social worldAalto University Doctoral Dissertations, 22/2018.
  9. Saarimäki, H. (2018). Decoding emotions from brain activity and connectivity patternsAalto University Doctoral Dissertations, 17/2018.
  10. Karlsson, H.K. (2016). Neuroreceptor availability and cerebral morphology in human obesity. Annales Universitatis Turkuensis D 1235. 
  11. Tuulari. J.J. (2015). Effects of obesity and weight loss following bariatric surgery on brain function, structural integrity and metabolism. Annales Universitatis Turkuensis D 1184.