{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,2]],"date-time":"2026-11-02T16:50:32Z","timestamp":1793638232440,"version":"4.5.1"},"reference-count":76,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,1,8]],"date-time":"2026-01-08T00:00:00Z","timestamp":1767830400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,2,12]],"date-time":"2026-02-12T00:00:00Z","timestamp":1770854400000},"content-version":"vor","delay-in-days":35,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/100000057","name":"U.S. Department of Health & Human Services | NIH | National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["P20GM121312"],"award-info":[{"award-number":["P20GM121312"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Commun Biol"],"DOI":"10.1038\/s42003-025-09498-3","type":"journal-article","created":{"date-parts":[[2026,1,8]],"date-time":"2026-01-08T20:34:27Z","timestamp":1767904467000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Age differences in neural discriminability of rumination and worry"],"prefix":"10.1038","volume":"9","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6829-4311","authenticated-orcid":false,"given":"Masaya","family":"Misaki","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aki","family":"Tsuchiyagaito","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Salvador M.","family":"Guinjoan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0825-3606","authenticated-orcid":false,"given":"Martin P.","family":"Paulus","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,1,8]]},"reference":[{"key":"9498_CR1","doi-asserted-by":"publisher","first-page":"400","DOI":"10.1111\/j.1745-6924.2008.00088.x","volume":"3","author":"S Nolen-Hoeksema","year":"2008","unstructured":"Nolen-Hoeksema, S., Wisco, B. E. & Lyubomirsky, S. Rethinking rumination. Perspect. Psychol. Sci. 3, 400\u2013424 (2008).","journal-title":"Perspect. Psychol. Sci."},{"key":"9498_CR2","doi-asserted-by":"publisher","first-page":"929","DOI":"10.1111\/bjc.12367","volume":"61","author":"AE Joubert","year":"2022","unstructured":"Joubert, A. E. et al. Understanding the experience of rumination and worry: a descriptive qualitative survey study. Br. J. Clin. Psychol. 61, 929\u2013946 (2022).","journal-title":"Br. J. Clin. Psychol."},{"key":"9498_CR3","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/0005-7967(83)90121-3","volume":"21","author":"TD Borkovec","year":"1983","unstructured":"Borkovec, T. D., Robinson, E., Pruzinsky, T. & DePree, J. A. Preliminary exploration of worry: some characteristics and processes. Behav. Res. Ther. 21, 9\u201316 (1983).","journal-title":"Behav. Res. Ther."},{"key":"9498_CR4","doi-asserted-by":"publisher","first-page":"218","DOI":"10.1037\/0003-066X.56.3.218","volume":"56","author":"BL Fredrickson","year":"2001","unstructured":"Fredrickson, B. L. The role of positive emotions in positive psychology: the broaden-and-build theory of positive emotions. Am. Psychol. 56, 218\u2013226 (2001).","journal-title":"Am. Psychol."},{"key":"9498_CR5","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1093\/geronb\/gbp132","volume":"65B","author":"S Scheibe","year":"2010","unstructured":"Scheibe, S. & Carstensen, L. L. Emotional aging: recent findings and future trends. J. Gerontol. B Psychol. Sci. Soc. Sci. 65B, 135\u2013144 (2010).","journal-title":"J. Gerontol. B Psychol. Sci. Soc. Sci."},{"key":"9498_CR6","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1037\/pspp0000453","volume":"125","author":"ST Charles","year":"2023","unstructured":"Charles, S. T. et al. Growing old and being old: emotional well-being across adulthood. J. Pers. Soc. Psychol. 125, 455\u2013469 (2023).","journal-title":"J. Pers. Soc. Psychol."},{"key":"9498_CR7","doi-asserted-by":"publisher","first-page":"e1019","DOI":"10.4088\/JCP.16m10671","volume":"77","author":"ML Thomas","year":"2016","unstructured":"Thomas, M. L. et al. Paradoxical trend for improvement in mental health with aging: a community-based study of 1,546 adults aged 21-100 years. J. Clin. Psychiatry 77, e1019\u2013e1025 (2016).","journal-title":"J. Clin. Psychiatry"},{"key":"9498_CR8","doi-asserted-by":"publisher","DOI":"10.1111\/nyas.15138","author":"M Mather","year":"2024","unstructured":"Mather, M. The emotion paradox in the aging body and brain. Ann. N. Y Acad. Sci. https:\/\/doi.org\/10.1111\/nyas.15138 (2024).","journal-title":"Ann. N. Y Acad. Sci."},{"key":"9498_CR9","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1037\/a0021285","volume":"26","author":"LL Carstensen","year":"2011","unstructured":"Carstensen, L. L. et al. Emotional experience improves with age: evidence based on over 10 years of experience sampling. Psychol. Aging 26, 21\u201333 (2011).","journal-title":"Psychol. Aging"},{"key":"9498_CR10","doi-asserted-by":"publisher","first-page":"734","DOI":"10.1037\/a0031180","volume":"104","author":"AJ Shallcross","year":"2013","unstructured":"Shallcross, A. J., Ford, B. Q., Floerke, V. & Mauss, I. B. Getting better with age: the relationship between age, acceptance, and negative affect. J. Pers. Soc. Psychol. 104, 734\u2013749 (2013).","journal-title":"J. Pers. Soc. Psychol."},{"key":"9498_CR11","doi-asserted-by":"publisher","first-page":"P313","DOI":"10.1093\/geronb\/62.6.P313","volume":"62","author":"JR Piazza","year":"2007","unstructured":"Piazza, J. R., Charles, S. T. & Almeida, D. M. Living with chronic health conditions: age differences in affective well-being. J. Gerontol.: Ser. B 62, P313\u2013P321 (2007).","journal-title":"J. Gerontol.: Ser. B"},{"key":"9498_CR12","doi-asserted-by":"publisher","first-page":"199","DOI":"10.1017\/S1041610219002084","volume":"32","author":"V Fernandez-Fernandez","year":"2020","unstructured":"Fernandez-Fernandez, V. et al. Emotion regulation processes as mediators of the impact of past life events on older adults\u2019 psychological distress. Int. Psychogeriatr. 32, 199\u2013209 (2020).","journal-title":"Int. Psychogeriatr."},{"key":"9498_CR13","doi-asserted-by":"publisher","first-page":"1063","DOI":"10.1080\/13607863.2015.1060944","volume":"20","author":"J Ricarte","year":"2016","unstructured":"Ricarte, J., Ros, L., Serrano, J. P., Martinez-Lorca, M. & Latorre, J. M. Age differences in rumination and autobiographical retrieval. Aging Ment. Health 20, 1063\u20131069 (2016).","journal-title":"Aging Ment. Health"},{"key":"9498_CR14","doi-asserted-by":"publisher","first-page":"80","DOI":"10.1093\/geronb\/gbz109","volume":"75","author":"L Emery","year":"2020","unstructured":"Emery, L., Sorrell, A. & Miles, C. Age differences in negative, but not positive, rumination. J. Gerontol. B Psychol. Sci. Soc. Sci. 75, 80\u201384 (2020).","journal-title":"J. Gerontol. B Psychol. Sci. Soc. Sci."},{"key":"9498_CR15","doi-asserted-by":"publisher","first-page":"192","DOI":"10.1521\/ijct.2008.1.3.192","volume":"1","author":"T Ehring","year":"2008","unstructured":"Ehring, T. & Watkins, E. R. Repetitive negative thinking as a transdiagnostic process. Int. J. Cogn. Ther. 1, 192\u2013205 (2008).","journal-title":"Int. J. Cogn. Ther."},{"key":"9498_CR16","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1002\/(SICI)1099-0879(199905)6:2<156::AID-CPP196>3.0.CO;2-A","volume":"6","author":"C Papageorgiou","year":"1999","unstructured":"Papageorgiou, C. & Wells, A. Process and meta-cognitive dimensions of depressive and anxious thoughts and relationships with emotional intensity. Clin. Psychol. Psychother. 6, 156\u2013162 (1999).","journal-title":"Clin. Psychol. Psychother."},{"key":"9498_CR17","doi-asserted-by":"publisher","first-page":"1577","DOI":"10.1016\/j.brat.2004.11.008","volume":"43","author":"E Watkins","year":"2005","unstructured":"Watkins, E., Moulds, M. & Mackintosh, B. Comparisons between rumination and worry in a non-clinical population. Behav. Res. Ther. 43, 1577\u20131585 (2005).","journal-title":"Behav. Res. Ther."},{"key":"9498_CR18","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1016\/j.beth.2006.03.003","volume":"38","author":"KA McLaughlin","year":"2007","unstructured":"McLaughlin, K. A., Borkovec, T. D. & Sibrava, N. J. The effects of worry and rumination on affect states and cognitive activity. Behav. Ther. 38, 23\u201338 (2007).","journal-title":"Behav. Ther."},{"key":"9498_CR19","doi-asserted-by":"publisher","first-page":"645","DOI":"10.1023\/B:COTR.0000045570.62733.9f","volume":"28","author":"GJ Siegle","year":"2004","unstructured":"Siegle, G. J., Moore, P. M. & Thase, M. E. Rumination: one construct, many features in healthy individuals, depressed individuals, and individuals with lupus. Cogn. Ther. Res. 28, 645\u2013668 (2004).","journal-title":"Cogn. Ther. Res."},{"key":"9498_CR20","doi-asserted-by":"publisher","first-page":"509","DOI":"10.1016\/j.janxdis.2010.03.008","volume":"24","author":"PM McEvoy","year":"2010","unstructured":"McEvoy, P. M., Mahoney, A. E. & Moulds, M. L. Are worry, rumination, and post-event processing one and the same? Development of the repetitive thinking questionnaire. J. Anxiety Disord. 24, 509\u2013519 (2010).","journal-title":"J. Anxiety Disord."},{"key":"9498_CR21","doi-asserted-by":"publisher","first-page":"926","DOI":"10.1177\/2167702614566603","volume":"3","author":"K Kircanski","year":"2015","unstructured":"Kircanski, K., Thompson, R. J., Sorenson, J., Sherdell, L. & Gotlib, I. H. Rumination and worry in daily life: examining the naturalistic validity of theoretical constructs. Clin. Psychol. Sci. 3, 926\u2013939 (2015).","journal-title":"Clin. Psychol. Sci."},{"key":"9498_CR22","doi-asserted-by":"publisher","first-page":"216","DOI":"10.1016\/j.jad.2018.08.037","volume":"241","author":"P Spinhoven","year":"2018","unstructured":"Spinhoven, P., van Hemert, A. M. & Penninx, B. W. Repetitive negative thinking as a predictor of depression and anxiety: a longitudinal cohort study. J. Affect. Disord. 241, 216\u2013225 (2018).","journal-title":"J. Affect. Disord."},{"key":"9498_CR23","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1017\/S0033291799001452","volume":"30","author":"AF Jorm","year":"2000","unstructured":"Jorm, A. F. Does old age reduce the risk of anxiety and depression? A review of epidemiological studies across the adult life span. Psychol. Med. 30, 11\u201322 (2000).","journal-title":"Psychol. Med."},{"key":"9498_CR24","doi-asserted-by":"publisher","first-page":"1188","DOI":"10.1093\/geront\/gnab116","volume":"61","author":"LL Carstensen","year":"2021","unstructured":"Carstensen, L. L. Socioemotional selectivity theory: the role of perceived endings in human motivation. Gerontologist 61, 1188\u20131196 (2021).","journal-title":"Gerontologist"},{"key":"9498_CR25","doi-asserted-by":"publisher","first-page":"380","DOI":"10.3389\/fpsyg.2014.00380","volume":"5","author":"U Kunzmann","year":"2014","unstructured":"Kunzmann, U., Kappes, C. & Wrosch, C. Emotional aging: a discrete emotions perspective. Front. Psychol. 5, 380 (2014).","journal-title":"Front. Psychol."},{"key":"9498_CR26","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1080\/10615806.2011.608126","volume":"25","author":"SC Segerstrom","year":"2012","unstructured":"Segerstrom, S. C. et al. Episodic repetitive thought: dimensions, correlates, and consequences. Anxiety Stress Coping 25, 3\u201321 (2012).","journal-title":"Anxiety Stress Coping"},{"key":"9498_CR27","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1080\/10615806.2014.947285","volume":"28","author":"SC Segerstrom","year":"2015","unstructured":"Segerstrom, S. C., Eisenlohr-Moul, T. A., Evans, D. R. & Ram, N. Repetitive thought dimensions, psychological well-being, and perceived growth in older adults: a multilevel, prospective study. Anxiety Stress Coping 28, 287\u2013302 (2015).","journal-title":"Anxiety Stress Coping"},{"key":"9498_CR28","doi-asserted-by":"publisher","first-page":"505","DOI":"10.1037\/a0019456","volume":"25","author":"SC Segerstrom","year":"2010","unstructured":"Segerstrom, S. C., Roach, A. R., Evans, D. R., Schipper, L. J. & Darville, A. K. The structure and health correlates of trait repetitive thought in older adults. Psychol. Aging 25, 505\u2013515 (2010).","journal-title":"Psychol. Aging"},{"key":"9498_CR29","doi-asserted-by":"publisher","first-page":"891","DOI":"10.1002\/jclp.20585","volume":"65","author":"SL Johnson","year":"2009","unstructured":"Johnson, S. L. & Jones, S. Cognitive correlates of mania risk: are responses to success, positive moods, and manic symptoms distinct or overlapping? J. Clin. Psychol. 65, 891\u2013905 (2009).","journal-title":"J. Clin. Psychol."},{"key":"9498_CR30","doi-asserted-by":"publisher","first-page":"1910","DOI":"10.1093\/cercor\/bhv001","volume":"26","author":"KA Lindquist","year":"2016","unstructured":"Lindquist, K. A., Satpute, A. B., Wager, T. D., Weber, J. & Barrett, L. F. The brain basis of positive and negative affect: evidence from a meta-analysis of the human neuroimaging literature. Cereb. Cortex 26, 1910\u20131922 (2016).","journal-title":"Cereb. Cortex"},{"key":"9498_CR31","doi-asserted-by":"publisher","DOI":"10.1016\/j.isci.2020.101060","volume":"23","author":"T Horikawa","year":"2020","unstructured":"Horikawa, T., Cowen, A. S., Keltner, D. & Kamitani, Y. The neural representation of visually evoked emotion is high-dimensional, categorical, and distributed across transmodal brain regions. iScience 23, 101060 (2020).","journal-title":"iScience"},{"key":"9498_CR32","doi-asserted-by":"publisher","first-page":"444","DOI":"10.1016\/j.tics.2016.03.011","volume":"20","author":"PA Kragel","year":"2016","unstructured":"Kragel, P. A. & LaBar, K. S. Decoding the nature of emotion in the brain. Trends Cogn. Sci. 20, 444\u2013455 (2016).","journal-title":"Trends Cogn. Sci."},{"key":"9498_CR33","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1016\/j.tics.2022.12.015","volume":"27","author":"C Westlin","year":"2023","unstructured":"Westlin, C. et al. Improving the study of brain-behavior relationships by revisiting basic assumptions. Trends Cogn. Sci. 27, 246\u2013257 (2023).","journal-title":"Trends Cogn. Sci."},{"key":"9498_CR34","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.abn8616","volume":"8","author":"B Kim Lux","year":"2022","unstructured":"Kim Lux, B., Andrews-Hanna, J. R., Han, J., Lee, E. & Woo, C.-W. When self comes to a wandering mind: brain representations and dynamics of self-generated concepts in spontaneous thought. Sci. Adv. 8, eabn8616 (2022).","journal-title":"Sci. Adv."},{"key":"9498_CR35","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1038\/s41586-022-05118-w","volume":"609","author":"AS Greene","year":"2022","unstructured":"Greene, A. S. et al. Brain-phenotype models fail for individuals who defy sample stereotypes. Nature 609, 109\u2013118 (2022).","journal-title":"Nature"},{"key":"9498_CR36","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1016\/j.neurobiolaging.2020.06.016","volume":"94","author":"AL Devitt","year":"2020","unstructured":"Devitt, A. L., Thakral, P. P., Szpunar, K., Addis, D. R. & Schacter, D. L. Age-related changes in repetition suppression of neural activity during emotional future simulation. Neurobiol. Aging 94, 287\u2013297 (2020).","journal-title":"Neurobiol. Aging"},{"key":"9498_CR37","doi-asserted-by":"publisher","first-page":"712","DOI":"10.1093\/scan\/nst035","volume":"9","author":"KK Szpunar","year":"2014","unstructured":"Szpunar, K. K., St Jacques, P. L., Robbins, C. A., Wig, G. S. & Schacter, D. L. Repetition-related reductions in neural activity reveal component processes of mental simulation. Soc. Cogn. Affect Neurosci. 9, 712\u2013722 (2014).","journal-title":"Soc. Cogn. Affect Neurosci."},{"key":"9498_CR38","unstructured":"Spielberger, C. D. Manual for the State-Trait Anxiety Inventory (Consulting Psychologists Press, 1970)."},{"key":"9498_CR39","doi-asserted-by":"publisher","DOI":"10.1016\/j.cpr.2020.101928","volume":"82","author":"KA Knowles","year":"2020","unstructured":"Knowles, K. A. & Olatunji, B. O. Specificity of trait anxiety in anxiety and depression: meta-analysis of the State-Trait Anxiety Inventory. Clin. Psychol. Rev. 82, 101928 (2020).","journal-title":"Clin. Psychol. Rev."},{"key":"9498_CR40","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1037\/0022-3514.61.1.115","volume":"61","author":"S Nolen-Hoeksema","year":"1991","unstructured":"Nolen-Hoeksema, S. & Morrow, J. A prospective study of depression and posttraumatic stress symptoms after a natural disaster: the 1989 Loma Prieta Earthquake. J. Pers. Soc. Psychol. 61, 115\u2013121 (1991).","journal-title":"J. Pers. Soc. Psychol."},{"key":"9498_CR41","doi-asserted-by":"publisher","first-page":"356","DOI":"10.1179\/2045772315Y.0000000024","volume":"38","author":"H Bertisch","year":"2015","unstructured":"Bertisch, H., Kalpakjian, C. Z., Kisala, P. A. & Tulsky, D. S. Measuring positive affect and well-being after spinal cord injury: development and psychometric characteristics of the SCI-QOL Positive Affect and Well-being bank and short form. J. Spinal Cord. Med. 38, 356\u2013365 (2015).","journal-title":"J. Spinal Cord. Med."},{"key":"9498_CR42","doi-asserted-by":"publisher","first-page":"487","DOI":"10.1016\/0005-7967(90)90135-6","volume":"28","author":"TJ Meyer","year":"1990","unstructured":"Meyer, T. J., Miller, M. L., Metzger, R. L. & Borkovec, T. D. Development and validation of the Penn State Worry Questionnaire. Behav. Res. Ther. 28, 487\u2013495 (1990).","journal-title":"Behav. Res. Ther."},{"key":"9498_CR43","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1016\/j.ijdevneu.2007.03.008","volume":"25","author":"NC McLaughlin","year":"2007","unstructured":"McLaughlin, N. C. et al. Diffusion tensor imaging of the corpus callosum: a cross-sectional study across the lifespan. Int. J. Dev. Neurosci. 25, 215\u2013221 (2007).","journal-title":"Int. J. Dev. Neurosci."},{"key":"9498_CR44","doi-asserted-by":"publisher","first-page":"352","DOI":"10.1177\/0963721410388395","volume":"19","author":"HL Urry","year":"2010","unstructured":"Urry, H. L. & Gross, J. J. Emotion regulation in older age. Curr. Dir. Psychol. Sci. 19, 352\u2013357 (2010).","journal-title":"Curr. Dir. Psychol. Sci."},{"key":"9498_CR45","doi-asserted-by":"publisher","first-page":"383","DOI":"10.1146\/annurev.psych.093008.100448","volume":"61","author":"ST Charles","year":"2010","unstructured":"Charles, S. T. & Carstensen, L. L. Social and emotional aging. Annu. Rev. Psychol. 61, 383\u2013409 (2010).","journal-title":"Annu. Rev. Psychol."},{"key":"9498_CR46","doi-asserted-by":"publisher","first-page":"587","DOI":"10.1007\/s00429-014-0803-z","volume":"220","author":"SC Krall","year":"2015","unstructured":"Krall, S. C. et al. The role of the right temporoparietal junction in attention and social interaction as revealed by ALE meta-analysis. Brain Struct. Funct. 220, 587\u2013604 (2015).","journal-title":"Brain Struct. Funct."},{"key":"9498_CR47","doi-asserted-by":"publisher","first-page":"796","DOI":"10.1002\/hbm.23068","volume":"37","author":"SC Krall","year":"2016","unstructured":"Krall, S. C. et al. The right temporoparietal junction in attention and social interaction: a transcranial magnetic stimulation study. Hum. Brain Mapp. 37, 796\u2013807 (2016).","journal-title":"Hum. Brain Mapp."},{"key":"9498_CR48","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1016\/j.neuropsychologia.2017.01.001","volume":"105","author":"KM Igelstrom","year":"2017","unstructured":"Igelstrom, K. M. & Graziano, M. S. A. The inferior parietal lobule and temporoparietal junction: a network perspective. Neuropsychologia 105, 70\u201383 (2017).","journal-title":"Neuropsychologia"},{"key":"9498_CR49","doi-asserted-by":"publisher","first-page":"1179","DOI":"10.1016\/j.biopsych.2004.02.023","volume":"55","author":"MP Paulus","year":"2004","unstructured":"Paulus, M. P., Feinstein, J. S., Simmons, A. & Stein, M. B. Anterior cingulate activation in high trait anxious subjects is related to altered error processing during decision making. Biol. Psychiatry 55, 1179\u20131187 (2004).","journal-title":"Biol. Psychiatry"},{"key":"9498_CR50","doi-asserted-by":"publisher","first-page":"1844","DOI":"10.1093\/brain\/aww069","volume":"139","author":"S Tolomeo","year":"2016","unstructured":"Tolomeo, S. et al. A causal role for the anterior mid-cingulate cortex in negative affect and cognitive control. Brain 139, 1844\u20131854 (2016).","journal-title":"Brain"},{"key":"9498_CR51","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms10904","volume":"7","author":"C Corradi-Dell\u2019Acqua","year":"2016","unstructured":"Corradi-Dell\u2019Acqua, C., Tusche, A., Vuilleumier, P. & Singer, T. Cross-modal representations of first-hand and vicarious pain, disgust and fairness in insular and cingulate cortex. Nat. Commun. 7, 10904 (2016).","journal-title":"Nat. Commun."},{"key":"9498_CR52","doi-asserted-by":"publisher","first-page":"331","DOI":"10.3389\/fpsyt.2019.00331","volume":"10","author":"JN Beadle","year":"2019","unstructured":"Beadle, J. N. & de la Vega, C. E. Impact of aging on empathy: review of psychological and neural mechanisms. Front. Psychiatry 10, 331 (2019).","journal-title":"Front. Psychiatry"},{"key":"9498_CR53","doi-asserted-by":"publisher","first-page":"1129","DOI":"10.1037\/a0025011","volume":"12","author":"JA Sze","year":"2012","unstructured":"Sze, J. A., Gyurak, A., Goodkind, M. S. & Levenson, R. W. Greater emotional empathy and prosocial behavior in late life. Emotion 12, 1129\u20131140 (2012).","journal-title":"Emotion"},{"key":"9498_CR54","doi-asserted-by":"publisher","DOI":"10.1016\/j.pnpbp.2021.110401","volume":"111","author":"JY Yun","year":"2021","unstructured":"Yun, J. Y. & Kim, Y. K. Graph theory approach for the structural-functional brain connectome of depression. Prog. Neuro-Psychopharmacol. Biol. Psychiatry 111, 110401 (2021).","journal-title":"Prog. Neuro-Psychopharmacol. Biol. Psychiatry"},{"key":"9498_CR55","doi-asserted-by":"publisher","first-page":"204","DOI":"10.1016\/j.pscychresns.2013.09.003","volume":"214","author":"W Truong","year":"2013","unstructured":"Truong, W. et al. Changes in cortical thickness across the lifespan in major depressive disorder. Psychiatry Res.: Neuroimaging 214, 204\u2013211 (2013).","journal-title":"Psychiatry Res.: Neuroimaging"},{"key":"9498_CR56","doi-asserted-by":"publisher","first-page":"619","DOI":"10.1136\/jnnp.2007.124651","volume":"79","author":"LL Herrmann","year":"2008","unstructured":"Herrmann, L. L., Le Masurier, M. & Ebmeier, K. P. White matter hyperintensities in late life depression: a systematic review. J. Neurol., Neurosurg. Psychiatry 79, 619\u2013624 (2008).","journal-title":"J. Neurol., Neurosurg. Psychiatry"},{"key":"9498_CR57","doi-asserted-by":"publisher","first-page":"488","DOI":"10.1192\/bjp.184.6.488","volume":"184","author":"AJ Lloyd","year":"2004","unstructured":"Lloyd, A. J. et al. Hippocampal volume change in depression: late- and early-onset illness compared. Br. J. Psychiatry 184, 488\u2013495 (2004).","journal-title":"Br. J. Psychiatry"},{"key":"9498_CR58","doi-asserted-by":"publisher","first-page":"954","DOI":"10.1038\/nbt.4200","volume":"36","author":"OG Sani","year":"2018","unstructured":"Sani, O. G. et al. Mood variations decoded from multi-site intracranial human brain activity. Nat. Biotechnol. 36, 954\u2013961 (2018).","journal-title":"Nat. Biotechnol."},{"key":"9498_CR59","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-20085-4","volume":"11","author":"H Kim","year":"2020","unstructured":"Kim, H., Smolker, H. R., Smith, L. L., Banich, M. T. & Lewis-Peacock, J. A. Changes to information in working memory depend on distinct removal operations. Nat. Commun. 11, 6239 (2020).","journal-title":"Nat. Commun."},{"key":"9498_CR60","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1016\/j.biopsycho.2011.05.003","volume":"87","author":"S Whittle","year":"2011","unstructured":"Whittle, S., Yucel, M., Yap, M. B. & Allen, N. B. Sex differences in the neural correlates of emotion: evidence from neuroimaging. Biol. Psychol. 87, 319\u2013333 (2011).","journal-title":"Biol. Psychol."},{"key":"9498_CR61","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1016\/S0006-3223(02)01866-8","volume":"54","author":"AJ Rush","year":"2003","unstructured":"Rush, A. J. et al. The 16-Item Quick Inventory of Depressive Symptomatology (QIDS), clinician rating (QIDS-C), and self-report (QIDS-SR): a psychometric evaluation in patients with chronic major depression. Biol. Psychiatry 54, 573\u2013583 (2003).","journal-title":"Biol. Psychiatry"},{"key":"9498_CR62","doi-asserted-by":"publisher","first-page":"447","DOI":"10.1007\/s10608-018-9901-1","volume":"42","author":"I Marchetti","year":"2018","unstructured":"Marchetti, I., Mor, N., Chiorri, C. & Koster, E. H. W. The Brief State Rumination Inventory (BSRI): validation and psychometric evaluation. Cogn. Ther. Res. 42, 447\u2013460 (2018).","journal-title":"Cogn. Ther. Res."},{"key":"9498_CR63","doi-asserted-by":"publisher","first-page":"385","DOI":"10.1016\/S0005-7967(03)00147-5","volume":"42","author":"A Wells","year":"2004","unstructured":"Wells, A. & Cartwright-Hatton, S. A short form of the metacognitions questionnaire: properties of the MCQ-30. Behav. Res. Ther. 42, 385\u2013396 (2004).","journal-title":"Behav. Res. Ther."},{"key":"9498_CR64","doi-asserted-by":"publisher","first-page":"1089","DOI":"10.1017\/S003329179900104X","volume":"29","author":"M Reynolds","year":"1999","unstructured":"Reynolds, M. & Wells, A. The Thought Control Questionnaire-psychometric properties in a clinical sample, and relationships with PTSD and depression. Psychol. Med. 29, 1089\u20131099 (1999).","journal-title":"Psychol. Med."},{"key":"9498_CR65","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1023\/B:JOBA.0000007455.08539.94","volume":"26","author":"KL Gratz","year":"2004","unstructured":"Gratz, K. L. & Roemer, L. Multidimensional assessment of emotion regulation and dysregulation: development, factor structure, and initial validation of the difficulties in emotion regulation scale. J. Psychopathol. Behav. Assess. 26, 41\u201354 (2004).","journal-title":"J. Psychopathol. Behav. Assess."},{"key":"9498_CR66","doi-asserted-by":"publisher","first-page":"1045","DOI":"10.1016\/j.paid.2006.04.010","volume":"41","author":"N Garnefski","year":"2006","unstructured":"Garnefski, N. & Kraaij, V. Cognitive emotion regulation questionnaire\u2014development of a short 18-item version (CERQ-short). Personal. Individ. Differ. 41, 1045\u20131053 (2006).","journal-title":"Personal. Individ. Differ."},{"key":"9498_CR67","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1002\/1522-2594(200007)44:1<162::AID-MRM23>3.0.CO;2-E","volume":"44","author":"GH Glover","year":"2000","unstructured":"Glover, G. H., Li, T. Q. & Ress, D. Image-based method for retrospective correction of physiological motion effects in fMRI: RETROICOR. Magn. Reson. Med. 44, 162\u2013167 (2000).","journal-title":"Magn. Reson. Med."},{"key":"9498_CR68","doi-asserted-by":"publisher","first-page":"644","DOI":"10.1016\/j.neuroimage.2007.11.059","volume":"40","author":"RM Birn","year":"2008","unstructured":"Birn, R. M., Smith, M. A., Jones, T. B. & Bandettini, P. A. The respiration response function: the temporal dynamics of fMRI signal fluctuations related to changes in respiration. Neuroimage 40, 644\u2013654 (2008).","journal-title":"Neuroimage"},{"key":"9498_CR69","doi-asserted-by":"publisher","first-page":"26","DOI":"10.1016\/j.media.2007.06.004","volume":"12","author":"BB Avants","year":"2008","unstructured":"Avants, B. B., Epstein, C. L., Grossman, M. & Gee, J. C. Symmetric diffeomorphic image registration with cross-correlation: evaluating automated labeling of elderly and neurodegenerative brain. Med. Image Anal. 12, 26\u201341 (2008).","journal-title":"Med. Image Anal."},{"key":"9498_CR70","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1016\/j.neuroimage.2010.04.246","volume":"52","author":"HJ Jo","year":"2010","unstructured":"Jo, H. J., Saad, Z. S., Simmons, W. K., Milbury, L. A. & Cox, R. W. Mapping sources of correlation in resting state FMRI, with artifact detection and removal. Neuroimage 52, 571\u2013582 (2010).","journal-title":"Neuroimage"},{"key":"9498_CR71","unstructured":"Feurer, M. et al. In Advances in Neural Information Processing Systems (eds Cortes, C. et al.) Vol. 28, 2962\u20142979 (Curran Associates, Inc., 2015)."},{"key":"9498_CR72","doi-asserted-by":"publisher","first-page":"2145","DOI":"10.1256\/003590002320603584","volume":"128","author":"SJ Mason","year":"2006","unstructured":"Mason, S. J. & Graham, N. E. Areas beneath the relative operating characteristics (ROC) and relative operating levels (ROL) curves: statistical significance and interpretation. Q. J. R. Meteorol. Soc. 128, 2145\u20132166 (2006).","journal-title":"Q. J. R. Meteorol. Soc."},{"key":"9498_CR73","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1111\/j.2517-6161.1995.tb02031.x","volume":"57","author":"Y Benjamini","year":"1995","unstructured":"Benjamini, Y. & Hochberg, Y. Controlling the false discovery rate: a practical and powerful approach to multiple testing. J. R. Stat. Soc. B 57, 289\u2013300 (1995).","journal-title":"J. R. Stat. Soc. B"},{"key":"9498_CR74","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18637\/jss.v067.i01","volume":"67","author":"D Bates","year":"2015","unstructured":"Bates, D., M\u00e4chler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1\u201348 (2015).","journal-title":"J. Stat. Softw."},{"key":"9498_CR75","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18637\/jss.v069.i01","volume":"69","author":"RV Lenth","year":"2016","unstructured":"Lenth, R. V. Least-squares means: the R package lsmeans. J. Stat. Softw. 69, 1\u201333 (2016).","journal-title":"J. Stat. Softw."},{"key":"9498_CR76","doi-asserted-by":"publisher","first-page":"152","DOI":"10.1089\/brain.2016.0475","volume":"7","author":"RW Cox","year":"2017","unstructured":"Cox, R. W., Chen, G., Glen, D. R., Reynolds, R. C. & Taylor, P. A. FMRI clustering in AFNI: false-positive rates redux. Brain Connect. 7, 152\u2013171 (2017).","journal-title":"Brain Connect."}],"container-title":["Communications Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-09498-3","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-09498-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-09498-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,12]],"date-time":"2026-02-12T13:04:27Z","timestamp":1770901467000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-09498-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,8]]},"references-count":76,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["9498"],"URL":"https:\/\/doi.org\/10.1038\/s42003-025-09498-3","relation":{},"ISSN":["2399-3642"],"issn-type":[{"value":"2399-3642","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,8]]},"assertion":[{"value":"20 February 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 December 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 January 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"223"}}