Aging Changes the Neural Correlates of Social Perspective Taking and Moral Judgements

Aging Changes the Neural Correlates of Social Perspective Taking and Moral Judgements | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Aging Changes the Neural Correlates of Social Perspective Taking and Moral Judgements Chenyi Chen, Yu-Chun Chen, Yang-Teng Fan, Róger Marcelo Martínez, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6859135/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Morality involves both cognitive and affective abilities, and while aging may hamper cognitive perspective-taking, it often enhances social-emotional focus. With strong motivation, older adults can perform as well as younger individuals in affective tasks. However, how aging affects the neural basis of Social Perspective Taking across these systems remains unclear. This study included young and older adults with varying implicit moral attitudes (mIAT) and Justice Sensitivity Inventory (JSI). fMRI scans were conducted while participants mentally simulated moral behaviors, such as helping or harming others. Older adults displayed higher mIAT D scores, reflecting greater conflict between positive valence and immoral actions. They also showed lower victim sensitivity on the JSI and attributed more praise and blame to moral actions, suggesting increased cognitive engagement in evaluating moral behaviors. Neuroimaging revealed that older adults showed reduced neural variation between helping and harming behaviors in the insula and anterior mid-cingulate cortex, indicating less engagement of affective systems. However, they maintained variation in the orbitofrontal and dorsolateral prefrontal cortex, relying more on cognitive systems. These findings suggest older adults compensate for diminished affective responses by engaging cognitive resources during moral evaluations and social perspective-taking, reflecting greater reliance on cognitive processing in differentiating moral behaviors. Biological sciences/Neuroscience/Cognitive neuroscience Biological sciences/Neuroscience/Emotion Biological sciences/Neuroscience/Social neuroscience Biological sciences/Psychology Biological sciences/Psychology/Human behaviour aging moral behavior moral attitudes social perspective-taking fMRI Figures Figure 1 Figure 2 Figure 3 Figure 4 INTRODUCTION Understanding how moral attitudes and social perspective-taking evolve with age is essential in a society where older adults increasingly contribute to caregiving and community leadership roles. For example, an elderly caregiver may confront complex decisions involving fairness and empathy daily—situations requiring both cognitive control and emotional sensitivity. These morally laden choices highlight the importance of probing how aging affects the mechanisms that underlie such decisions. Defining Perspective-Taking in Moral Cognition Perspective-taking enables individuals to infer others' thoughts and emotions and is a cornerstone of moral cognition. It includes cognitive elements (understanding mental states) and affective elements (empathizing with emotions), which recruit overlapping yet distinct neural systems—prefrontal cortices for cognitive and limbic/ventromedial prefrontal areas for affective components (Healey and Grossman, 2018 ). Findings on aging and perspective-taking are mixed. Older adults show declines in lab-based affective tasks (Fernandes et al., 2019a ; Pratt et al., 1996a ), yet perform comparably to younger adults when motivated (Zhang et al., 2013 ). They tend to use more positive language in narrative retellings (Sullivan et al., 2010 ) and excel in social inference despite visual/spatial declines (De Lillo and Ferguson, 2023 ). Perspective-taking deficits in aging have been linked to reduced dorsomedial prefrontal cortex activity (Moran et al., 2012 ). The dynamic integration theory (Labouvie-Vief, 2003 ) posits that emotional and cognitive complexity peaks in midlife and declines in older adulthood, giving way to emotional prioritization (O'Brien et al., 2013 ). Age-related declines in executive function, memory, and processing speed (Hambrick et al., 1999 ; Salthouse, 1994 ; West, 1996 ) may underlie diminished cognitive perspective-taking (Fernandes et al., 2019b ; Maylor et al., 2002 ; Pratt et al., 1996b ), while moral reasoning shifts from intent to outcome (Margoni et al., 2018 ). Socioemotional Aging and Moral Reasoning Despite cognitive declines, older adults often show enhanced emotional regulation and social focus (Isaacowitz et al., 2021 ), extending to moral domains (Margoni et al., 2023 ). They report greater relational satisfaction (Diener et al., 1999 ; Lang and Carstensen, 1994 ; Litwin, 2001 ), demonstrate stronger memory for moral events (Narvaez et al., 2011 ), and favor deontological reasoning linked to moral idealism and affective sensitivity (Arutyunova et al., 2016a ; McNair et al., 2018 ). This paradox—cognitive decline alongside social-emotional effectiveness—may reflect task-context differences (Daley et al., 2023 ; Marsiske et al., 1995; Spreng and Turner, 2019 ). Findings remain mixed: older adults rate immoral acts as more wrong but show reduced emotional arousal (Lu and Fung, 2019 ). Electrophysiology suggests diminished emotional engagement (Fernandes et al., 2019a ), while neuroimaging shows reduced affective but preserved cognitive empathy circuits in aging (Chen et al., 2014a ). Moral evaluations involve both cognitive and affective processes across brain regions including the amygdala, insula, anterior mid-cingulate cortex (aMCC), temporal pole, right temporoparietal junction (rTPJ), orbitofrontal cortex (OFC), ventromedial and dorsolateral prefrontal cortex (dlPFC), medial prefrontal cortex (mPFC), and striatum (Buckholtz and Marois, 2012 ; Moll et al., 2002 ). These systems support moral cognition through arousal, valuation, and social reasoning (Decety et al., 2012 ; Pascual et al., 2013 ). Older adults tend to favor deontological choices, particularly when the decision feels intuitive, but show no age-related difference when utilitarian responses are also intuitive. This bias is associated with enhanced connectivity between the posterior medial core of the default mode network (pmDN) and the dorsal attention network, alongside reduced segregation of the pmDN from the broader brain network (Huang et al., 2021 ). Stronger DMN connectivity supported better memory across ages and reduced utilitarian choices in younger adults. In older adults, similar moral choices involved greater DMN–Salience Network interaction, suggesting age-related shifts in how DMN supports memory and moral decisions (Daley and Kensinger, 2022 ). Implicit and Explicit Moral Judgments Across the Lifespan Explicit moral judgments are deliberate and reflect rational processing grounded in values and norms, while implicit judgments are automatic and emotionally driven (Cameron et al., 2017 ; Greenwald et al., 1998). Aging may alter this balance, with older adults possibly leaning more on affective or habitual responses in moral contexts. Explicit attitudes are commonly assessed via tools like the Justice Sensitivity Inventory (Schmitt et al., 2010 ), and implicit attitudes through the Implicit Association Test (IAT). Given age-related slowing, we applied a z-score transformation to the IAT to account for processing speed differences (Faust et al., 1999 ; Hummert et al., 2002 ). Together, distinctions between cognitive and affective processing, perspective-taking, and implicit versus explicit judgment styles help illuminate age-related changes in moral reasoning. While behavioral studies highlight decision speed and bias shifts, fMRI enables examination of the neural underpinnings. In this study, we used fMRI with a button-trigger, first-person simulation of moral scenarios to explore these mechanisms across age groups (Chen et al., 2022 ; Ruby and Decety, 2001 ). Based on current understanding of age-related shifts in cognitive and affective processing—characterized by increased socio-affective motivation and decreased cognitive ability—we hypothesize the following: (1) First-Person Perspective-Taking Reaction Times: Due to the high socio-affective motivation tied to moral and immoral agency (Zhang et al., 2013 ), older adults will perform comparably to younger adults. (2) Moral Evaluations of Observed Actions: Older adults may exhibit heightened praise/blame evaluations of moral/immoral behaviors, consistent with findings of increased sensitivity to victims’ unpleasantness during evaluations of harmful actions (Chen et al. ( 2014c )). (3) Implicit and Explicit Moral Attitudes: Implicit attitudes, closely tied to automatic affective inclinations (Isaacowitz et al., 2021 ), may increase with age, whereas explicit attitudes, associated with deliberative cognitive evaluation (Chen et al., 2014b; Spreng and Turner, 2019 ), are expected to remain stable or decline in older adults compared to younger adults. (4) fMRI BOLD Responses: We hypothesize reduced activation in affective brain regions such as the anterior insular cortex (AIC) and anterior mid-cingulate cortex (aMCC) related to pain empathy in older adults, with stable activity in the medial prefrontal cortex and posterior superior temporal sulcus associated with cognitive empathy and perceived agency (Chen et al., 2014b). This shift may indicate more efficient valence processing in older adults, accompanied by increased reliance on cognitive brain subsystems and heightened moral sensitivity due to changes in the cognitive-affective balance. MATERIALS AND METHODS Participants The study enrolled two groups of right-handed, ethnic Chinese participants from the community: (1) 31 young adults (15 males, aged 20 to 31 years, mean ± standard deviation (SD): 23 ± 2.2); and (2) 32 older adults (11 males, 65 to 80, 69 ± 4.0). One older subject was excluded due to excessive head movement during fMRI scanning, leaving N = 62 for the fMRI data analysis. All participants were screened to ensure that they had no history of neurological or psychiatric disorders, had normal or corrected-normal visual acuity, and were taking no medications at the time of testing. Older participants were further screened for medication use and recent surgical procedures. This study's experimental protocols received ethical approval from the Institutional Review Board of National Yang-Ming University (Protocol Number: YM102033). All participants gave written informed consent for the study, which was approved by local ethic committee, and was conducted in accordance with the Declaration of Helsinki . Visual Stimuli Forty-five validated animations from previous fMRI studies were presented to participants (Chen et al., 2020a ; Chen et al., 2020b ; Cheng et al., 2021a ; Decety and Porges, 2011 ). Each animation comprised three images with duration of 1000, 200, and 1000 milliseconds, respectively, and which portrayed the following scenarios: (1) a person who is taking an action to physically harming one another person (harming); and (2) a person who is alleviating physical pain from a suffering person (helping). One additional baseline stimuli depicted people carrying out an action that was irrelevant to the other one (neutral). Procedures Participants performed moral judgments on moral dilemmas and completed the dispositional Justice Sensitivity Inventory (JSI) and the morally-laden implicit association test (mIAT) before fMRI scanning (supplementary Figure s1 ). During fMRI scanning, we made use of mental simulations of social interactions, and a button-trigger design was adopted with reference to the current literature (Chen et al., 2022 ). Participants first imagined themselves in the scenario and pressed a handheld button when ready for the action to proceed. They viewed an initial image of the action, then pressed the button to trigger the following two images. There were two runs—one with simulated harming and the other with helping actions—intermixed with neutral action blocks. Before each run, participants were instructed to mentally simulate themselves as the agent of either harming or helping actions from a first-person perspective. Participants could press the button at their own pace (mean reaction time ± standard error: young adults, 1238 ± 172 ms; older adults, 1798 ± 170 ms), which initiated the complete action sequence. The task aimed to measure the time participants needed to mentally prepare for simulating the actions. Quicker responses suggested easier engagement and lower cognitive load. Before the formal trials, participants practiced with three trials and feedback to ensure understanding. The button-press was designed to help immerse participants in the virtual agent's role. After fMRI scanning, the visual stimuli were presented, and participants, with the use of a computer-based seven-point visual analogue scale, were asked to evaluate how much praise they attributed to themselves as the imagined protagonist in the helping actions or how much blame they attributed to themselves as the imagined protagonist in the harming actions. Dispositional justice sensitivity (JSI) The JSI is a self-reported measure that assesses four aspects of justice sensitivity, with scores ranging from 1 (strongly disagree) to 6 (strongly agree). Three perspectives—observer, beneficiary, and perpetrator—are often combined into a single "other-oriented sensitivity" score (Edele et al., 2013 ; Gollwitzer et al., 2009 ). While the JSI measured participants’ sensitivity to injustice, unfairness, or immoral scenarios, self-oriented sensitivity referred to situations in which participants themselves were the victims of injustice, making the sensitivity directed toward themselves. In contrast, other-oriented sensitivity referred to sensitivity toward injustice experienced by others (See supplementary materials for detailed methods). Utilitarian judgments on moral dilemmas Based on previous work (Chen et al., 2016 ; Greene et al., 2009 ; Greene et al., 2004 ; Greene et al., 2001 ; Huebner et al., 2011 ) ( http://www.cell.com/neuron/supplemental/S0896-6273(04)00634-8 ), forty-eight moral dilemmas were selected to create two versions of the moral judgment task balanced for emotional intensity (Koenigs et al., 2007 ). Each version included 12 dilemmas: six personal (e.g., pushing a stranger in the trolley dilemma) and six impersonal (e.g., flipping a switch). Personal dilemmas involved direct harm, while impersonal involved indirect harm. Personal dilemmas were further classified as involving inevitable or evitable harm, with higher moral permissibility judgments for inevitable harm. The dilemmas were translated from English to Chinese and back-translated for accuracy. Participants read and responded at their own pace. Implicit moral attitudes (mIAT) The mIAT was modified with animations and words for the stimuli. Verbal stimuli included 26 extremely pleasant and 26 extremely unpleasant words, selected from high-frequency Chinese words (Chen et al., 2020a ; Chen et al., 2022 ; Chen et al., 2020b ; Martínez et al., 2020 ). The mIAT performance, as indexed by D scores, represents implicit moral attitudes. The D score was calculated by subtracting the mean RT of congruent (immoral-negative) blocks from that of incongruent (immoral-positive) blocks and dividing it by the pooled SD across the two blocks (Nosek et al., 2014 ). Higher D scores might come from higher RTs for incongruent blocks or lower RTs for congruent blocks (See supplementary materials for detailed methods). fMRI acquisition, data processing and analysis Scanning employed a mixed design (19.1 s ON/13.2 ± 4.4 s OFF) with two runs, each consisting of six ON blocks (two each of harming, helping, and neutral scenarios) and six OFF blocks. ON blocks included five trials, each lasting 2200 ms, with pseudo-randomized sequence presentation. Scanning was performed using a 3T Siemens Magnetom Trio-Tim MRI, and BOLD fMRI signals were collected. Functional and structural images were processed using SPM12 and analyzed in MATLAB. Functional images were realigned, co-registered, normalized to MNI space, and smoothed. A voxel-by-voxel regression analysis modeled three conditions (harming, helping, and neutral) to capture neuro-hemodynamic responses, with contrasts for harming vs. neutral and helping vs. neutral. These first-level contrast images were entered into an ANOVA to examine age and scenario effects, with whole-brain activation corrected for multiple comparisons (FWE rate p < 0.05). To assess how aging moderated neural responses, ROI data from cognitive and affective networks were extracted based on predefined coordinates from the literature and prior studies. Four cognitive ROIs (dlPFC, dmPFC, TPJ, OFC) and four affective ROIs (amygdala, insula, aMCC, temporal pole) were analyzed using MarsBaR in SPM12 (Chen et al., 2022 ; Cheng et al., 2021b ; Moran et al., 2012 ) (See supplementary materials for detailed methods). RESULTS Behavioral Performance Demographics Table 1 lists demographic data. Although younger participants scored higher on the Mini-Mental Status Examination (MMSE) (young: 29.8 ± 0.1; old: 28.8 ± 0.2, t = 4.49, p < 0.001), none of the older participants were suffering from dementia, as indicated by all MMSE scores being higher than 25 (ranging from 27 to 30) (Folstein et al., 1975 ). No subjects had any evidence of depression, as indicated by scores lower than 5 on the Chinese version of the Geriatric Depression Scale (young: 2.13 ± 0.37; old: 1.47 ± 0.25, t = 1.47, p = 0.15) (Mui, 1996 ). Table 1 Demographic and dispositional variables of the participants. Young Old ( N = 31) ( N = 32) Mean SE Mean SE p Age (years) 23.03 0.4 69.28 0.7 < 0.001 Gender (male%) 48 9.1 34 8.5 0.27 MMSE 29.77 0.09 28.78 0.2 < 0.001 Depression 2.13 0.37 1.47 0.25 0.15 Education (years) 16.97 0.33 14.32 0.55 < 0.001 mIAT D-score 0.61 0.05 0.91 0.06 0.001 JSI Self-oriented 3.70 0.18 2.87 0.21 0.004 Other-oriented 4.46 0.15 4.41 0.18 0.84 JSI total score 4.27 0.11 4.02 0.16 0.21 Moral dilemmas (endorsement %) Non-moral 63.08 5.86 56.6 4.79 0.39 Impersonal 52.26 7.11 60.0 7.18 0.45 Inevitable 52.42 6.49 46.88 6.12 0.54 Evitable 20.43 2.28 25.52 2.9 0.17 Moral evaluation Blame 5.08 0.19 6 0.26 0.006 Praise 4.7 0.22 6.3 0.18 < 0.001 Abbreviations: SE, standard error; MMSE, Mini-Mental Status Examination; mIAT, morality implicit association test; JSI, Justice Sensitivity Inventory; Depression, Chinese version of the Geriatric Depression Scale. Age differences in implicit moral attitudes For implicit moral attitudes, older adults exhibited higher D scores than young adults (young: 0.61 ± 0.05; old: 0.94 ± 0.06, t = -3.59, p = 0.001) (Fig. 1 A). Because the IAT relies on RTs, higher D scores found in older adults not only reflected implicit attitudes but also age differences in processing speeds. Given that older adults exhibited a greater increase in reaction times (RTs) compared to younger adults, it is essential to disentangle the effect of implicit attitudes from the general age-related slowing. Accordingly, trial-level z-score transformations were applied separately to congruent and incongruent trials (Hummert et al., 2002 ). Resulting z-score transformation latencies were then entered into an ANOVA: 2 (age: young vs. old) × 2 (congruency: congruent vs. incongruent). Results showed a main effect of congruency ( F 1, 61 = 343.51, p < 0.001, pη 2 = 0.85; congruent vs. incongruent: -0.38 ± 0.02 vs. 0.39 ± 0.02, mean ± SE) and an interaction effect between age and congruency ( F 1, 61 = 15.46, p < 0.001, pη 2 = 0.20), while the main effect of age ( F 1, 61 = 1.85, p = 0.18) did not reached significance. Post-hoc tests revealed that, when compared to young adults, older adults showed significant shorter latencies in the congruent trials (young vs. old: -0.3 ± 0.027 vs. -0.46 ± 0.03 s, t = 3.84, p < 0.001), but longer latencies in the incongruent trials (young vs. old: 0.31 ± 0.027 vs. 0.48 ± 0.032 s, t = -3.99, p < 0.001). Age differences in dispositional justice sensitivity The 2 (age: young vs. old; a between-subject factor) × 2 (perspective: self-oriented vs. other-oriented; a within-subject factor) mixed ANOVA on JSI scores showed a main effect of age ( F 1, 61 = 5.25, p = 0.025, pη 2 = 0.079; young vs. old: 4.08 ± 0.14 vs. 3.64 ± 0.14), perspective ( F 1, 61 = 44.99, p < 0.001, pη 2 = 0.424; self- vs. other-oriented: 3.28 ± 0.14 vs. 4.44 ± 0.12), and an interaction effect between age and perspective ( F 1, 61 = 5.3, p = 0.025, pη 2 = 0.08). Post-hoc tests revealed that, when compared to young adults, older adults showed significantly reduced victim sensitivity toward self-oriented concerns (young vs. old: 3.7 ± 0.18 vs. 2.87 ± 0.21, t = 2.95, p = 0.004), but comparable sensitivities toward other-oriented concerns (young vs. old: 4.46 ± 0.15 vs. 4.41 ± 0.18, t = 0.21, p = 0.84) (Fig. 1 B). Age differences in moral evaluations Due to the RT data not being normally distributed, the RTs were base-10 log-transformed for further analyses. A 2 (age: young vs. old) × 3 (scenario: harming vs. helping vs. neutral) repeated ANOVA on log-transformed RTs revealed a main effect of scenario ( F 2, 122 = 51.11, p < 0.001, pη 2 = 0.46) with the longest RTs for harming behavior (3.19 ± 0.03 s) as compared to helping (3.07 ± 0.03 s, t = 9.52, p < 0.001) and neutral actions (3.03 ± 0.03 s, t = 8.41, p < 0.001). There was neither a main effect of age ( F 1, 61 = 1.26, p = 0.27) nor an age × scenario interaction ( F 1, 61 = 0.01, p = 0.92) (Fig. 2 A). Subjective ratings included praise for interpersonal assistance and blame for harming behaviors. Compared to younger participants, older adults attributed more praise when helping others (young vs. old: 4.7 ± 0.22 vs. 6.3 ± 0.18, t = -5.72, p < 0.001), and more blame when harming others (young vs. old: 5.1 ± 0.19 vs. 6.0 ± 0.26, t = -2.85, p = 0.006) (Fig. 2 B). Age differences in moral dilemmas The 2 (age: young vs. old; a between-subject factor) × 4 (dilemma type: nonmoral, impersonal, personal-evitable, or personal-inevitable; a within-subject factor) mixed ANOVA on the endorsement of utilitarian decisions produced a main effect of dilemma type ( F 3, 183 = 20.33, p < 0.001, pη 2 = 0.25), while there was neither an effect of age ( F 1, 61 = 0.002, p = 0.97) nor age × dilemma type interaction ( F 3, 183 = 0.97, p = 0.36). The evitable harm (22.98 ± 1.85) was judged as less morally endorsed than did the dilemmas with nonmoral (59.84 ± 3.78, t = -10.19, p < 0.001), impersonal (56.13 ± 5.05, t = -7.2, p < 0.001), and the inevitable harm (49.65 ± 4.46, t = -5.75, p < 0.001). Neuroimaging results Neuro-hemodynamic response to moral scenarios The entire sample of 62 participants (regardless of age) showed significant neuro-hemodynamic increases in the network of regions involved in both cognitive and affective processing of moral brain ( k > 20, p < 0.05, FWE-corrected) (Healey and Grossman, 2018 ). Regions with greater activity in response to scenarios involving interpersonal harm vs. assistance included the dmPFC, dlPFC, TPJ, OFC, insula, aMCC, temporal pole, hippocampus, caudate, amygdala, postcentral gyrus, and occipital cortex. The reverse contrast showed no significant cluster (supplementary Table s1 ). ROI results Regarding the age effect on neuro-hemodynamic responses, while there was no survival cluster of the main effect of age (young vs. old) found in the voxel-wise analyses, the interaction between age (young vs. old) and scenario (harming vs. helping) was identified among several ROIs, including the dlPFC ( F 1, 60 = 4.1, p = 0.047, pη 2 = 0.06), OFC ( F 1, 60 = 4.77, p = 0.033, pη 2 = 0.07), insula ( F 1, 60 = 4.14, p = 0.046, pη 2 = 0.07), and aMCC ( F 1, 60 = 5.83, p = 0.019, pη 2 = 0.09) (supplementary Table s2). Post-hoc analyses indicated that scenario effects in the dlPFC, OFC, insula, and aMCC showed opposite directions depending on the factor of age. Harming compared to helping actions showed increased activity in the aMCC (harming vs. helping: 0.45 ± 0.11 vs. -0.05 ± 0.1, t = 2.85, p = 0.008) with a trend in the insula (0.25 ± 0.09 vs. -0.04 ± 0.1, t = 1.75, p = 0.09) in young adults, but none in older adults (insula: -0.03 ± 0.07 vs. 0.06 ± 0.07, t = -1.04, p = 0.306; aMCC: 0.07 ± 0.08 vs. 0.08 ± 0.07, t = -0.06, p = 0.955). On the other hand, harming as compared to helping actions showed increased activities in the dlPFC (0.2 ± 0.12 vs. -0.15 ± 0.09, t = 2.16, p = 0.039) and OFC (0.38 ± 0.19 vs. -0.22 ± 0.19, t = 2.43, P = 0.021) in older adults, but none in young adults (dlPFC: -0.002 ± 0.13 vs. 0.15 ± 0.1, t = -0.82, p = 0.418; OFC: 0.03 ± 0.2 vs. 0.32 ± 0.18, t = -0.89, p = 0.38) (Fig. 3 ). Regarding brain-behavior correlations, the young and older groups showed a double dissociation between neural correlates and behavioral assessments (Fig. 4 ). When adopting harming actions, the amygdala activity positively predicted D scores of implicit moral attitudes in young adults ( r = 0.43, p = 0.015), but not in older adults ( r = -0.25, p = 0.18). The rTPJ activity negatively predicted the self-related justice sensitivity within the young (r= -0.502, P = 0.004), but not within the elders ( r = 0.29, P = 0.11). The temporal pole activity positively predicted the blame ratings in young adults ( r = 0.8, p = 0.038), but not in older adults ( r = -0.03, p = 0.87). Fisher’s z tests confirmed the significant dissociations in the amygdala (z = 2.68, p = 0.007) and rTPJ (z = 3.18, p = 0.002). On the other hand, adopting helping actions had no such dissociation. DISCUSSION This study revealed age-related changes in implicit and explicit moral attitudes and their neural correlates during perspective-taking of moral behavior. While aging had no effect on moral dilemmas, older adults exhibited higher D scores on the mIAT and lower victim sensitivity on the JSI. Reaction times for social perspective-taking showed no age-related differences, but older adults attributed greater praise and blame to moral behaviors. Reduced differential BOLD activity was observed in the insula and anterior mid-cingulate cortex (aMCC) for helping versus harming, while the orbitofrontal cortex (OFC) and dorsolateral prefrontal cortex (dlPFC) maintained differentiation. Unlike younger adults, correlations between BOLD activity and moral attitudes were absent in older adults. These findings suggest greater moral cognitive evaluation in older adults, reflecting a neural shift in cognitive and affective processing with aging. Contrary to expectations, moral dilemmas showed no age effects or interactions, potentially due to cultural differences. Increased deontological judgments in older adults are more prominent in Western cultures (Arutyunova et al., 2016a ). Chinese individuals are less likely to sacrifice one to save five (Gold et al., 2014 ), judging avoidable harm more harshly than impersonal or inevitable harm. Cultural differences between East and West influence moral concepts. In collectivist Chinese culture, morality emphasizes social harmony and respect for hierarchy, extending to behaviors deemed impolite in Western contexts, such as not offering a seat (Nichols, 2022). By contrast, Western individualistic cultures prioritize utilitarian ethics, favoring decisions maximizing collective welfare, even with trade-offs (Arutyunova et al., 2016b; Bentahila et al., 2021 ; Hong and Sciences, 2023). These cultural norms may explain why age-related shifts in moral reasoning observed in Western contexts are less pronounced in Chinese culture, where Confucian values persist across age groups. Older adults showed significantly higher D scores on the implicit moral attitudes, as indicated by the mIAT, even after controlling for age-related slowing effects on RTs. The higher D scores might have resulted from either higher RTs for incongruent (immoral-positive) blocks or lower RTs for congruent (immoral-negative) blocks (Nosek et al., 2014 ). The D scores on the mIAT were predictive of actor-observer bias when attributing blame for immoral behaviors and individual differences during moral computation (Chen et al., 2020a ; Chen et al., 2022 ; Chen et al., 2020b ). Higher D scores, revealing increased conflict between positive/negative valence and immoral/moral behaviors, might indicate that older adults experienced more difficulty in distinguishing the greater cognitive evaluation triggered by moral behaviors from the conflict arising from word valence pairs. In parallel, they also explicitly attributed more praise and blame to their own moral behaviors. Additionally, older adults reported lower victim sensitivity toward self-oriented concerns but comparable sensitivity toward other-oriented concerns on the JSI. This is supported by previous findings indicating that older adults tend to exhibit more morally idealistic beliefs (Margoni et al., 2018 ). These behavioral results support our hypotheses: (1) Older adults, driven by high socio-affective motivation related to moral and immoral agency, demonstrated performance comparable to younger adults, with no significant differences observed in reaction times. (2) Consistent with previous findings, older adults exhibited increased sensitivity, assigning heightened praise or blame to moral and immoral behaviors relative to younger adults. (3) Older adults showed stronger implicit moral attitudes, indicating greater conflict between automatic affective inclinations toward positive or negative valence and moral or immoral actions. Interestingly, on explicit measures of justice sensitivity, older adults showed reduced sensitivity when they themselves were the victims of injustice, reflecting diminished self-oriented sensitivity, while maintaining sensitivity toward injustices experienced by others. This pattern may be partially attributed to high socio-affective motivation and the cultural framework discussed above. In contrast to young adults, older adults retained differential BOLD activities in the OFC and dlPFC in response to harming vs. helping others, but exhibited reduced differential activities in the insula and aMCC. When perceiving moral scenarios, older relative to younger adults showed an intent-to-outcome shift specific to harming actions, which was associated with a decline in theory of mind abilities (Margoni et al., 2018 ). In line with age-related changes in empathy for pain, activity in the anterior insula and anterior mid-cingulate cortex (aMCC) exhibited age-related declines, whereas activity in the medial prefrontal cortex (mPFC) and right temporoparietal junction (rTPJ) remained preserved in older adults, even as they reported increased unpleasantness (Chen et al., 2014c ). Considering the heightened behavioral sensitivity and reduced activity in the anterior insular cortex (AIC) and anterior mid-cingulate cortex (aMCC) associated with the affective components of pain empathy and affective perspective-taking in older adults, these findings—alongside increased activity observed in the orbitofrontal cortex (OFC) and right temporoparietal junction (rTPJ)—may indicate a shifting balance between cognitive and affective processing with age (Healey and Grossman, 2018 ). This shift suggests that valence processing within the affective brain system may become more efficient in older adults, leading to reduced activation in these regions. Consequently, older adults may allocate more resources to cognitive brain subsystems, which typically show age-related declines (Brehmer et al., 2012 ; Kensinger et al., 2017 ). Correlations between BOLD activities and moral attitudes and ratings found in young adults did not exist in older adults. Based on the same paradigm, young adults revealed that the OFC mediated the mIAT–RTs association, and the rTPJ mediated the JSI–warm-glow linkage (Chen et al., 2022 ). Young adults also displayed positive correlations of the mIAT with neural responses in the OFC and temporal poles to actor-observer asymmetry (Chen et al., 2020a ). Such brain-behavior correlations vanished with aging. One previous fMRI study revealed that older adults frequently exhibited greater magnitudes and extents of activation of the BOLD signals compared to younger adults. This additional activation might reflect compensatory recruitment associated with functional and structural deterioration of neural resources (Woodard and Sugarman, 2012 ). In interpreting our findings, it is essential to consider the nature of the moral task used in this study. While participants were instructed to mentally simulate themselves as agents in morally laden actions, the button-trigger design with sequential image presentation places the task at an intersection between first-person and third-person moral judgment paradigms. Prior work has shown that first-person tasks tend to elicit greater emotional engagement and activation in affective networks (Greene et al., 2009 ), whereas third-person tasks are more likely to involve reflective perspective-taking and recruit cognitive control regions (Koenigs et al., 2007 ). Our paradigm, with its explicit instruction for self-agency but lack of real-world consequence, may have evoked a hybrid engagement, consistent with findings that moral cognition involves both intuitive and deliberative processes (Daley and Kensinger, 2022 ; Huang et al., 2021 ). Age-related neural differences observed here—specifically, the diminished insula and aMCC response in older adults alongside preserved OFC and dlPFC activity—may reflect a shift in the balance of these networks, supporting the idea that aging modulates the integration of affective and cognitive moral systems(Daley and Kensinger, 2022 ). These distinctions underscore the importance of carefully considering task framing in moral neuroscience research, particularly when comparing across age groups. A few limitations of the current work should be clarified for future research. For instance, we adopted a first-person perspective, rather than a third-person perspective, thereby limiting direct comparisons with previous available literatures. First- or third-person perspective in moral cognition involves distinct neural processes (Avram et al., 2014 ; Chen et al., 2020a ). In young adults, dlPFC and rTPJ activities were involved in the asymmetry between first-hand experiencing and passive viewing moral behaviors, and the first-person relative to third-person perspective elicited stronger activity in the aMCC (Chen et al., 2020a ). To discern any aging impact on the neural underpinnings of moral behavior, it would be better to use a first-person perspective. Finally, our sample included only young and older adults, but did not include middle-aged adults. This might not be an optimal design, and future studies are warranted with longitudinal research in clinical populations or large community-based samples. All in all, this study demonstrated that aging has an impact on the neural underpinnings of moral attitudes and behaviors. The results suggest a reduced engagement of neural affective subsystems, with older adults relying more on the neural cognitive subsystems of the prefrontal cortex. These findings could contribute to the existing body of developmental research (Cowell and Decety, 2015 ; Decety et al., 2012 ) in demonstrating the neuro-cognitive trajectories of morality over a person’s lifespan. Given the fact that morality arguably serves adaptive social functions during one’s lifespan, especially in adulthood as individuals become responsible for their own and for others well-being (Gebremariam and Sadana, 2019 ), the present findings may contribute to a better understanding of healthy aging, and help provide insights into neurodegenerative disorders with socio-emotional deficits. Declarations ACKNOWLEDGMENTS We thank Jwu-Jiun Chang for assisting with data collection and preparation of stimuli. We thank Prof. Jean Decety for the morally-laden scenarios used as the stimuli; without his generous contribution such an endeavor would not have been possible. FUNDINGS The study was funded by the National Science and Technology Council, R.O.C. (NSTC 113-2410-HA49-065; 113-2410-H-038-034 -; 113-2410-H-028 -005 -), National Yang Ming Chiao Tung University Hospital (RD2025-002), and the Higher Education Sprout Project by the Ministry of Education (MOE) in Taiwan (TMU-114-N-05), Taiwan. CONFLICT OF INTEREST None of the authors has any conflicts of interest to declare. DATA AVAILABILITY STATEMENT The data presented in this study are available on request from the corresponding author. CRediT authorship contribution statement C.C. and Y.C. conceived and designed the study. C.C. and Y.C. supervised the study. C.C. and Y.C. wrote the main manuscript text. Y.C.C. curated the data. C.C. formally analyzed the data. C.C., Y.C.C., Y.T.F., R.M.M., and Y.C. reviewed and edited the manuscript. Declaration of Generative AI and AI-Assisted Technologies: The author(s) acknowledge the use of generative AI and AI-assisted technologies during the preparation of this manuscript. Specifically, OpenAI’s ChatGPT was used to improve the clarity, grammar, and readability of the English language in the writing process. No content was generated or interpreted by the AI; all conceptual, analytical, and interpretative work was conducted by the authors. The authors take full responsibility for the integrity and accuracy of the content presented. References Arutyunova, K.R., Alexandrov, Y.I., Hauser, M.D., 2016a. Sociocultural Influences on Moral Judgments: East-West, Male-Female, and Young-Old. Front Psychol 7, 1334. Arutyunova, K.R., Alexandrov, Y.I., Hauser, M.D.J.F.i.p., 2016b. Sociocultural influences on moral judgments: East–west, male–female, and young–old. 7, 1334. Avram, M., Hennig-Fast, K., Bao, Y., Poppel, E., Reiser, M., Blautzik, J., Giordano, J., Gutyrchik, E., 2014. Neural correlates of moral judgments in first- and third-person perspectives: implications for neuroethics and beyond. BMC Neurosci 15, 39. Bentahila, L., Fontaine, R., Pennequin, V.J.F.i.P., 2021. Universality and cultural diversity in moral reasoning and judgment. 12, 764360. Brehmer, Y., Westerberg, H., Bäckman, L.J.F.i.h.n., 2012. Working-memory training in younger and older adults: training gains, transfer, and maintenance. 6, 63. Buckholtz, J.W., Marois, R., 2012. The roots of modern justice: cognitive and neural foundations of social norms and their enforcement. Nat Neurosci 15, 655-661. Cameron, C.D., Payne, B.K., Sinnott-Armstrong, W., Scheffer, J.A., Inzlicht, M., 2017. Implicit moral evaluations: A multinomial modeling approach. Cognition 158, 224-241. Chen, C., Decety, J., Huang, P.C., Chen, C.Y., Cheng, Y., 2016. Testosterone administration in females modulates moral judgment and patterns of brain activation and functional connectivity. Human Brain Mapping 37, 3417-3430. Chen, C., Martinez, R.M., Chen, Y., Cheng, Y., 2020a. Pointing fingers at others: The neural correlates of actor-observer asymmetry in blame attribution. Neuropsychologia 136, 107281. Chen, C., Martinez, R.M., Chen, Y.C., Fan, Y.T., Cheng, Y., 2022. The neural mediators of moral attitudes and behaviors. Behav Brain Res 430, 113934. Chen, C., Martinez, R.M., Cheng, Y., 2020b. The key to group fitness: The presence of another synchronizes moral attitudes and neural responses during moral decision-making. Neuroimage 213, 116732. Chen, Y.-C., Chen, C.-C., Decety, J., Cheng, Y., 2014a. Aging is associated with changes in the neural circuits underlying empathy. Neurobiology of aging 35, 827-836. Chen, Y.-C., Chen, C.-C., Decety, J., Cheng, Y.J.N.o.a., 2014b. Aging is associated with changes in the neural circuits underlying empathy. 35, 827-836. Chen, Y.C., Chen, C.C., Decety, J., Cheng, Y., 2014c. Aging is associated with changes in the neural circuits underlying empathy. Neurobiol Aging 35, 827-836. Cheng, Y., Chou, J., Martinez, R.M., Fan, Y.T., Chen, C., 2021a. Psychopathic traits mediate guilt-related anterior midcingulate activity under authority pressure. Sci Rep 11, 14856. Cheng, Y., Chou, J., Martínez, R.M., Fan, Y.-T., Chen, C., 2021b. Psychopathic traits mediate guilt-related anterior midcingulate activity under authority pressure. Scientific reports 11, 1-11. Cowell, J.M., Decety, J., 2015. The neuroscience of implicit moral evaluation and its relation to generosity in early childhood. Curr Biol 25, 93-97. Daley, R.T., Kensinger, E.A.J.A., Neuropsychology,, Cognition, 2023. Cognitive decline, socioemotional change, or both? How the science of aging can inform future research on sacrificial moral dilemmas. 30, 272-299. Daley, R.T., Kensinger, E.A.J.N., 2022. Age-related differences in Default Mode Network resting-state functional connectivity but not gray matter volume relate to sacrificial moral decision-making and working memory performance. 177, 108399. De Lillo, M., Ferguson, H.J.J.J.o.E.P.G., 2023. Perspective-taking and social inferences in adolescents, young adults, and older adults. 152, 1420. Decety, J., Michalska, K.J., Kinzler, K.D., 2012. The contribution of emotion and cognition to moral sensitivity: a neurodevelopmental study. Cereb Cortex 22, 209-220. Decety, J., Porges, E.C., 2011. Imagining being the agent of actions that carry different moral consequences: an fMRI study. Neuropsychologia 49, 2994-3001. Diener, E., Suh, E.M., Lucas, R.E., Smith, H.L., 1999. Subjective well-being: Three decades of progress. Psychological Bulletin 125, 276-302. Edele, A., Dziobek, I., Keller, M., 2013. Explaining altruistic sharing in the dictator game: the role of affective empathy, cognitive empathy, and justice sensitivity. Learning and Individual Differences 24. Faust, M.E., Balota, D.A., Spieler, D.H., Ferraro, F.R.J.P.b., 1999. Individual differences in information-processing rate and amount: implications for group differences in response latency. 125, 777. Fernandes, C., Gonçalves, A., Pasion, R., Ferreira-Santos, F., Barbosa, F., Martins, I., Marques-Teixeira, J., 2019a. Age-related decline in emotional perspective-taking: Its effect on the late positive potential. Cognitive, Affective, Behavioral Neuroscience 19, 109-122. Fernandes, C., Goncalves, A.R., Pasion, R., Ferreira-Santos, F., Barbosa, F., Martins, I.P., Marques-Teixeira, J., 2019b. Age-related decline in emotional perspective-taking: Its effect on the late positive potential. Cogn Affect Behav Neurosci 19, 109-122. Folstein, M.F., Folstein, S.E., McHugh, P.R., 1975. "Mini-mental state". A practical method for grading the cognitive state of patients for the clinician. J Psychiatr Res 12, 189-198. Gebremariam, K.M., Sadana, R., 2019. On the ethics of healthy ageing: setting impermissible trade-offs relating to the health and well-being of older adults on the path to universal health coverage. Int J Equity Health 18, 140. Gold, N., Colman, A.M., Pulford, B.D., 2014. Cultural differences in responses to real-life and hypothetical trolley problems. Judgment and Decision Making 9, 65-76. Gollwitzer, M., Rothmund, T., Pfeiffer, A., Ensenbach, C., 2009. Why and when justice sensitivty leads to pro- and antisocial behavior. Journal of Research in Personality 43, 999-1005. Greene, J.D., Cushman, F.A., Stewart, L.E., Lowenberg, K., Nystrom, L.E., Cohen, J.D., 2009. Pushing moral buttons: The interaction between personal force and intention in moral judgment. Cognition 111, 364-371. Greene, J.D., Nystrom, L.E., Engell, A.D., Darley, J.M., Cohen, J.D., 2004. The neural bases of cognitive conflict and control in moral judgment. Neuron 44, 389-400. Greene, J.D., Sommerville, R.B., Nystrom, L.E., Darley, J.M., Cohen, J.D., 2001. An fMRI investigation of emotional engagement in moral judgment. Science 293, 2105-2108. Greenwald, A.G., McGhee, D.E., Schwartz, J.L.J.J.o.p., psychology, s., 1998. Measuring individual differences in implicit cognition: the implicit association test. 74, 1464. Hambrick, D.Z., Salthouse, T.A., Meinz, E.J., 1999. Predictors of crossword puzzle proficiency and moderators of age-cognition relations. J Exp Psychol Gen 128, 131-164. Healey, M.L., Grossman, M., 2018. Cognitive and affective perspective-taking: evidence for shared and dissociable anatomical substrates. Frontiers in neurology 9, 491. Hong, H.J.J.o.E., Humanities, Sciences, S., 2023. Cultural Differences in Moral Judgement. 10, 146-151. Huang, S., Faul, L., Sevinc, G., Mwilambwe-Tshilobo, L., Setton, R., Lockrow, A.W., Ebner, N.C., Turner, G.R., Spreng, R.N., De Brigard, F.J.P., aging, 2021. Age differences in intuitive moral decision-making: Associations with inter-network neural connectivity. 36, 902. Huebner, B., Hauser, M.D., Pettit, P., 2011. How the source, inevitability and means of bringing about harm interact in folk‐moral judgments. Mind Language 26, 210-233. Hummert, M.L., Garstka, T.A., O'Brien, L.T., Greenwald, A.G., Mellott, D.S., 2002. Using the implicit association test to measure age differences in implicit social cognitions. Psychol Aging 17, 482-495. Isaacowitz, D.M., Freund, A.M., Mayr, U., Rothermund, K., Tobler, P.N.J.T.J.o.G.S.B., 2021. Age-related changes in the role of social motivation: Implications for healthy aging. 76, S115-S124. Kensinger, E.A., Gutchess, A.H.J.J.o.G.S.B.P.S., Sciences, S., 2017. Cognitive aging in a social and affective context: Advances over the past 50 years. 72, 61-70. Koenigs, M., Young, L., Adolphs, R., Tranel, D., Cushman, F., Hauser, M., Damasio, A., 2007. Damage to the prefrontal cortex increases utilitarian moral judgements. Nature 446, 908-911. Labouvie-Vief, G., 2003. Dynamic Integration: Affect, Cognition, and the Self in Adulthood. Current Directions in Psychological Science 12, 201-206. Lang, F.R., Carstensen, L.L., 1994. Close emotional relationships in late life: further support for proactive aging in the social domain. Psychol Aging 9, 315-324. Litwin, H., 2001. Social network type and morale in old age. Gerontologist 41, 516-524. Lu, M., Fung, H.H., 2019. Moral gain or decay? examing age-related changes in moral judgment. Innov Aging 3, S785. Margoni, F., Geipel, J., Hadjichristidis, C., Bakiaj, R., Surian, L.J.C.s., 2023. Age‐Related Differences in Moral Judgment: The Role of Probability Judgments. 47, e13345. Margoni, F., Geipel, J., Hadjichristidis, C., Surian, L., 2018. Moral Judgment in Old Age. Exp Psychol 65, 105-114. Marsiske, M., Willis, S.L.J.P., aging, 1995. Dimensionality of everyday problem solving in older adults. 10, 269. Martínez, R.M., Chen, C.-Y., Liao, T.-T., Cheng, Y., Fan, Y.-T., Chou, S.-H., Chen, C.J.F.i.P., 2020. The multifaceted effects of serotonin transporter polymorphism (5-HTTLPR) on anxiety, implicit moral attitudes, and harmful behaviors. 11, 1521. Maylor, E.A., Moulson, J.M., Muncer, A.M., Taylor, L.A., 2002. Does performance on theory of mind tasks decline in old age? Br J Psychol 93, 465-485. McNair, S., Okan, Y., Hadjichristidis, C., de Bruin, W.B., 2018. Age differences in moral judgment: older adults are more deontological than younger adults. Behavioral Decsion Making 32, 47-60. Moll, J., de Oliveira-Souza, R., Bramati, I.E., Grafman, J., 2002. Functional networks in emotional moral and nonmoral social judgments. Neuroimage 16, 696-703. Moran, J.M., Jolly, E., Mitchell, J.P., 2012. Social-cognitive deficits in normal aging. J Neurosci 32, 5553-5561. Mui, A.C., 1996. Geriatric Depression Scale as a community screening instrument for elderly Chinese immigrants. Int Psychogeriatr 8, 445-458. Narvaez, D., Radvansky, G.A., Lynchard, N.A., Copeland, D.E., 2011. Are older adults more attuned to morally charged information? Exp Aging Res 37, 398-434. Nosek, B.A., Bar-Anan, Y., Sriram, N., Axt, J., Greenwald, A.G., 2014. Understanding and using the brief Implicit Association Test: recommended scoring procedures. PLoS One 9, e110938. O'Brien, E., Konrath, S.H., Grühn, D., Hagen, A.L., 2013. Empathic concern and perspective taking: linear and quadratic effects of age across the adult life span. J Gerontol B Psychol Sci Soc Sci 68, 168-175. Pascual, L., Rodrigues, P., Gallardo-Pujol, D., 2013. How does morality work in the brain? A functional and structural perspective of moral behavior. Front Integr Neurosci 7, 65. Pratt, M.W., Diessner, R., Pratt, A., Hunsberger, B., Pancer, S.M., 1996a. Moral and social reasoning and perspective taking in later life: a longitudinal study. Psychology Aging 11, 66. Pratt, M.W., Diessner, R., Pratt, A., Hunsberger, B., Pancer, S.M., 1996b. Moral and social reasoning and perspective taking in later life: a longitudinal study. Psychol Aging 11, 66-73. Ruby, P., Decety, J., 2001. Effect of subjective perspective taking during simulation of action: a PET investigation of agency. Nat Neurosci 4, 546-550. Salthouse, T.A., 1994. Aging associations: influence of speed on adult age differences in associative learning. J Exp Psychol Learn Mem Cogn 20, 1486-1503. Schmitt, M., Baumert, A., Gollwitzer, M., Maes, J., 2010. The Justice Sensitivity Inventory: Factorial validity, location in the personality facet space, demorgraphic pattern, and normative data. Social Justice Research 23, 211-238. Spreng, R.N., Turner, G.R.J.P.o.P.S., 2019. The shifting architecture of cognition and brain function in older adulthood. 14, 523-542. Sullivan, S.J., Mikels, J.A., Carstensen, L.L.J.P., aging, 2010. You never lose the ages you’ve been: Affective perspective taking in older adults. 25, 229. West, R.L., 1996. An application of prefrontal cortex function theory to cognitive aging. Psychol Bull 120, 272-292. Woodard, J.L., Sugarman, M.A., 2012. Functional Magnetic Resonance Imaging in Aging and Dementia: Detection of Age-Related Cognitive Changes and Prediction of Cognitive Decline, in: Pardon, M.-C., Bondi, M.W. (Eds.), Behavioral Neurobiology of Aging. Springer Berlin Heidelberg, Berlin, Heidelberg, pp. 113-136. Zhang, X., Fung, H.H., Stanley, J.T., Isaacowitz, D.M., Ho, M.Y.J.D.P., 2013. Perspective taking in older age revisited: a motivational perspective. 49, 1848. Additional Declarations No competing interests reported. Supplementary Files mIATagesupplementary.docx Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6859135","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":521231210,"identity":"a27f118a-c75d-4d89-86da-11ce49ad3f98","order_by":0,"name":"Chenyi Chen","email":"","orcid":"","institution":"Taipei Medical University","correspondingAuthor":false,"prefix":"","firstName":"Chenyi","middleName":"","lastName":"Chen","suffix":""},{"id":521231211,"identity":"c96eca4e-fdf1-4b02-ac3d-af2bb12309f5","order_by":1,"name":"Yu-Chun Chen","email":"","orcid":"","institution":"National Taiwan University of Sport","correspondingAuthor":false,"prefix":"","firstName":"Yu-Chun","middleName":"","lastName":"Chen","suffix":""},{"id":521231212,"identity":"329a6445-6e14-415a-9b6f-3ecb823c4221","order_by":2,"name":"Yang-Teng Fan","email":"","orcid":"","institution":"Yuan Ze University","correspondingAuthor":false,"prefix":"","firstName":"Yang-Teng","middleName":"","lastName":"Fan","suffix":""},{"id":521231213,"identity":"873cfa6a-20e7-4568-a01a-7999ec45c4d6","order_by":3,"name":"Róger Marcelo Martínez","email":"","orcid":"","institution":"Taipei Medical University","correspondingAuthor":false,"prefix":"","firstName":"Róger","middleName":"Marcelo","lastName":"Martínez","suffix":""},{"id":521231214,"identity":"295c44ec-bc04-4f25-acda-07374cfa25b1","order_by":4,"name":"Yawei Cheng","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA0klEQVRIiWNgGAWjYHACNhAhB+UwE6/FGKGFjUgtiQ1EazG4kfzswccdtenbJZKfSTBUWCc2yPcYENCSZm4488zx3J0z0swkGM6kJzaw8RDQcjuHTZq37Vjuhhs5bBKMbYeBWng3ENbyt+1YugFYyz9itTC21SRAtDQQoUXy/jMzyd62A4Ybzjwztkg4lm7cxpb/Aa8WvjOHn0n8bKuTNzie/PDGhxpr2X7mYwl4tSgcAFOHGRgEgApBagnGpHwDmKpjYOA/QEjtKBgFo2AUjFQAAB1qR+hBFnsZAAAAAElFTkSuQmCC","orcid":"","institution":"National Yang Ming Chiao Tung University Hospital","correspondingAuthor":true,"prefix":"","firstName":"Yawei","middleName":"","lastName":"Cheng","suffix":""}],"badges":[],"createdAt":"2025-06-10 05:08:24","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6859135/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6859135/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":92681612,"identity":"8498bd1f-19c1-4f3d-984e-4da0afb1741c","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"doc","order_by":0,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":335872,"visible":true,"origin":"","legend":"","description":"","filename":"mIATagemx06132.doc","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/b54b1f8741fc0aa7344dd94a.doc"},{"id":92682538,"identity":"627234b6-3370-4807-942d-3e15da903d72","added_by":"auto","created_at":"2025-10-03 01:17:55","extension":"tif","order_by":1,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":696908,"visible":true,"origin":"","legend":"","description":"","filename":"Fig1.tif","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/b6a91c862b0d1654a2afbec7.tif"},{"id":92682539,"identity":"5b949a84-2c87-450d-ac1a-d961c675b93c","added_by":"auto","created_at":"2025-10-03 01:17:55","extension":"tif","order_by":2,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":835264,"visible":true,"origin":"","legend":"","description":"","filename":"Fig2.tif","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/4efc6ec522cfbfe886049dc7.tif"},{"id":92681616,"identity":"1f512058-c7d1-4055-b791-5bfddccb87de","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"tif","order_by":3,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1713928,"visible":true,"origin":"","legend":"","description":"","filename":"Fig3.tif","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/05d43dd672ef205fe1e570c7.tif"},{"id":92681619,"identity":"24c4981b-468b-4478-8d72-719a5aa2d6cf","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"tif","order_by":4,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1384560,"visible":true,"origin":"","legend":"","description":"","filename":"Fig4.tif","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/75d0a3fcdcf6ef26aba1e142.tif"},{"id":92681615,"identity":"2c354f5f-1bd9-4b24-86f0-9ad32a565b37","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"json","order_by":5,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":7241,"visible":true,"origin":"","legend":"","description":"","filename":"ce84dd5bfbab43d2b752d27761af533a.json","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/d51088241d66a342d086c6ec.json"},{"id":92681625,"identity":"a6d1c2fb-e7db-494f-a170-a01a4ac594a7","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"docx","order_by":6,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":289214,"visible":true,"origin":"","legend":"","description":"","filename":"mIATagesupplementary.docx","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/ff8e97ac0f1350c2d0a2ebf6.docx"},{"id":92682965,"identity":"4925c0b6-e39d-4bd2-9e2d-d6c4a71f5218","added_by":"auto","created_at":"2025-10-03 01:25:55","extension":"xml","order_by":7,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":158848,"visible":true,"origin":"","legend":"","description":"","filename":"ce84dd5bfbab43d2b752d27761af533a1enriched.xml","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/430e9f55fb781a3d19a8e253.xml"},{"id":92681626,"identity":"2a5f0c24-55d2-4738-941b-56675165c69f","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"tif","order_by":8,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":696908,"visible":true,"origin":"","legend":"","description":"","filename":"Fig1.tif","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/2937d3d25ea48438eb4e0305.tif"},{"id":92681621,"identity":"1c457b4a-0b88-4852-8f8d-87869a426ff4","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"tif","order_by":9,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":835264,"visible":true,"origin":"","legend":"","description":"","filename":"Fig2.tif","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/025e78b31410401ec836b119.tif"},{"id":92682540,"identity":"47357456-ee13-4e19-bf14-23521643be40","added_by":"auto","created_at":"2025-10-03 01:17:55","extension":"tif","order_by":10,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1713928,"visible":true,"origin":"","legend":"","description":"","filename":"Fig3.tif","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/0970822b6254591aec1aa68d.tif"},{"id":92681631,"identity":"e153fd83-d95e-4e3f-96e7-694e76fdc368","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"tif","order_by":11,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1384560,"visible":true,"origin":"","legend":"","description":"","filename":"Fig4.tif","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/b6b847c24bd2beeb938a3131.tif"},{"id":92681630,"identity":"cca40c4a-51b3-4e69-b63d-0f1d6e16d6cd","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"png","order_by":12,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":133375,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineFig1.png","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/cdf5af873766bec40e029f36.png"},{"id":92681620,"identity":"d2b4d549-4777-480e-87d6-43b8b41fe91b","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"png","order_by":13,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":149901,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineFig2.png","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/7df22d8451a06d03473b7ba0.png"},{"id":92681623,"identity":"bbfea172-c89b-4abe-94cd-337a9ee83c3e","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"png","order_by":14,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":250353,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineFig3.png","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/cfa5729771d665cccbb295cf.png"},{"id":92682542,"identity":"769a5e3d-b696-4c88-91e7-14cf1ab3c546","added_by":"auto","created_at":"2025-10-03 01:17:55","extension":"png","order_by":15,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":190144,"visible":true,"origin":"","legend":"","description":"","filename":"OnlineFig4.png","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/a66a35bbfac5984084ba8767.png"},{"id":92681633,"identity":"a271b584-ce0b-4a45-af12-33dfe896d06b","added_by":"auto","created_at":"2025-10-03 01:09:56","extension":"xml","order_by":16,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":155270,"visible":true,"origin":"","legend":"","description":"","filename":"ce84dd5bfbab43d2b752d27761af533a1structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/26e045d021e492dc89933e63.xml"},{"id":92681629,"identity":"e0d6da20-1911-4c83-ab66-80224b5b24a9","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"html","order_by":17,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":171630,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/6a08ecbccd16bfa713aae39e.html"},{"id":92681609,"identity":"47b39a0a-602e-4fec-9bf9-9919351e2f07","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1061834,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eAging changes in theimplicit moral attitude (mIAT) and Justice Sensitivity Inventory (JSI).\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e(A)\u003c/strong\u003e. Older adults exhibited higher D scores on the mIAT than younger adults (young: 0.61±0.05; old: 0.94±0.06,\u003cem\u003e p\u003c/em\u003e = 0.001). A 2 (age: young vs. old) ´ 2 (congruency: congruent vs. incongruent) ANOVA on the z-score transformation latencies (reaction times (RTs)) revealed an interaction. Post-hoc tests showed that compared to younger adults, older adults showed significant shorter latencies in the congruent trials (young vs. old: -0.3±0.027 vs. -0.46±0.03 s, \u003cem\u003ep \u003c/em\u003e\u0026lt; 0.001), but longer latencies in the incongruent trials (young vs. old: 0.31±0.027 vs. 0.48±0.032 s, \u003cem\u003et\u003c/em\u003e = -3.99, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e(B)\u003c/strong\u003eThe 2 (age: young vs. old) ´ 2 (perspective: self-oriented vs. other-oriented) mixed ANOVA on JSI scores showed a significant interaction between age and perspective. Post-hoc tests revealed that compared to younger adults, older adults showed significantly reduced victim sensitivity toward self-related concerns (young vs. old: 3.7±0.18 vs. 2.87±0.21, \u003cem\u003ep\u003c/em\u003e = 0.004), but comparable sensitivity toward other-related concerns (young vs. old: 4.46 ± 0.15 vs. 4.41 ± 0.18, \u003cem\u003ep\u003c/em\u003e = 0.84).\u003c/p\u003e","description":"","filename":"Fig1.png","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/7d4eea97e093d9ed964f03a6.png"},{"id":92681611,"identity":"ebf276f4-a77e-4c06-ae07-e1851ed32f62","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1256254,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eAging changes in the subjective ratings and reaction times (RTs) of moral behavior.\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e(A)\u003c/strong\u003e. A 2 (age: young vs. old) ´ 3 (scenario: harming vs. helping vs. neutral) repeated ANOVA on log-transformed RTs revealed a main effect of scenario with the longest RTs for harming behavior (3.19±0.03) as compared to helping (3.07±0.03, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001) and neutral actions (3.03±0.03, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001). There was neither an age effect nor an age ´ scenario interaction for RTs of moral behavior.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e(B).\u003c/strong\u003e Subjective ratings included praise for interpersonal assistance, and blame for harming behaviors. Compared to younger adults, older adults attributed more praise when helping others (young vs. old: 4.7 ± 0.22 vs. 6.3 ± 0.18, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001), and more blame when harming others (young vs. old: 5.08 ± 0.19 vs. 6 ± 0.26, \u003cem\u003ep\u003c/em\u003e = 0.006).\u003c/p\u003e","description":"","filename":"Fig2.png","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/6c6daf00e018e7f3921d61ef.png"},{"id":92681624,"identity":"09d55eeb-22cd-480d-8aaa-44110958e772","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":3397285,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eHemodynamic responses to moral behaviors in young and old adults.\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e(A).\u003c/strong\u003e The entire sample of 62 participants (regardless of age) showed significant neuro-hemodynamic increases in the network of regions involved in both cognitive and affective processing (k \u0026gt; 20, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05, family-wise error (FWE) corrected). Regions with greater activity in response to scenarios involving interpersonal harm vs. interpersonal assistance included the dmPFC, dlPFC, TPJ, OFC, insula, aMCC, temporal pole, hippocampus, caudate, amygdala, postcentral gyrus, and occipital cortex.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e(B).\u003c/strong\u003e ROI analyses identified several ROIs showing a significant interaction between the factor age (young vs. old) and scenario (harming vs. helping), including the dlPFC, OFC, insula, and aMCC. Harming actions compared to helping actions showed increased activity in affective processing by the aMCC (harming vs. helping: 0.45±0.11 vs. -0.05±0.1, \u003cem\u003ep\u003c/em\u003e = 0.008) and insula (0.25±0.09 vs. -0.04±0.1, \u003cem\u003ep\u003c/em\u003e = 0.09, \u003cem\u003emarginal significance\u003c/em\u003e) in younger adults but none in the older adults (all \u003cem\u003ep\u003c/em\u003e \u0026gt; 0.3). On the other hand, harming actions compared to helping actions showed increased activity in the cognitive processing by the dlPFC (0.2±0.12 vs. -0.15±0.09, \u003cem\u003ep\u003c/em\u003e = 0.039) and OFC (0.38±0.19 vs. -0.22±0.19, \u003cem\u003ep \u003c/em\u003e= 0.021) in older adults, but none in younger adults (all \u003cem\u003ep\u003c/em\u003e \u0026gt; 0.3)\u003c/p\u003e","description":"","filename":"Fig3.png","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/5827eb02fa9e09d5f543c1e2.png"},{"id":92681622,"identity":"5dc8c266-4352-4156-8d10-574e6546713b","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":2857437,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eDouble dissociation between the neuro-hemodynamic responses and behavioral assessments in young and old adults.\u003c/strong\u003e When adopting harming behavior, the amygdala activity positively predicted implicit moral attitudes of D-scores in younger adults (\u003cem\u003er\u003c/em\u003e = 0.43, \u003cem\u003ep \u003c/em\u003e= 0.015), but not in older adults (\u003cem\u003er\u003c/em\u003e = -0.25, \u003cem\u003ep \u003c/em\u003e= 0.178). Activity in the rTPJ negatively predicted the self-related justice sensitivity in younger adults (\u003cem\u003er\u003c/em\u003e = -0.50, \u003cem\u003ep \u003c/em\u003e= 0.004), but not in older adults (\u003cem\u003er\u003c/em\u003e = 0.29, \u003cem\u003ep \u003c/em\u003e= 0.114). Activity in the temporal pole positively predicted blame ratings in younger adults (\u003cem\u003er \u003c/em\u003e= 0.375, \u003cem\u003ep \u003c/em\u003e= 0.038), but not in older adults (\u003cem\u003er\u003c/em\u003e = -0.03, \u003cem\u003ep \u003c/em\u003e= 0.872). Fisher’s z tests confirmed significant dissociations in the amygdala (z = 2.68, \u003cem\u003ep \u003c/em\u003e= 0.007) and rTPJ (z = 3.18, \u003cem\u003ep \u003c/em\u003e= 0.002).\u003c/p\u003e","description":"","filename":"Fig4.png","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/b01ac3cd2091a61852be9fb0.png"},{"id":95527274,"identity":"6dfd5232-d6bd-48c7-b170-f9c7b7fe2e5c","added_by":"auto","created_at":"2025-11-10 10:12:37","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":10098208,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/7c53b8b3-e300-4080-a196-afa5941b5858.pdf"},{"id":92681614,"identity":"81152b73-7156-4785-8a28-bc18cadeb834","added_by":"auto","created_at":"2025-10-03 01:09:55","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":289214,"visible":true,"origin":"","legend":"","description":"","filename":"mIATagesupplementary.docx","url":"https://assets-eu.researchsquare.com/files/rs-6859135/v1/6b715179c5dd8c7df4fdf575.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Aging Changes the Neural Correlates of Social Perspective Taking and Moral Judgements","fulltext":[{"header":"INTRODUCTION","content":"\u003cp\u003eUnderstanding how moral attitudes and social perspective-taking evolve with age is essential in a society where older adults increasingly contribute to caregiving and community leadership roles. For example, an elderly caregiver may confront complex decisions involving fairness and empathy daily\u0026mdash;situations requiring both cognitive control and emotional sensitivity. These morally laden choices highlight the importance of probing how aging affects the mechanisms that underlie such decisions.\u003c/p\u003e\n\u003ch3\u003eDefining Perspective-Taking in Moral Cognition\u003c/h3\u003e\n\u003cp\u003ePerspective-taking enables individuals to infer others' thoughts and emotions and is a cornerstone of moral cognition. It includes cognitive elements (understanding mental states) and affective elements (empathizing with emotions), which recruit overlapping yet distinct neural systems\u0026mdash;prefrontal cortices for cognitive and limbic/ventromedial prefrontal areas for affective components (Healey and Grossman, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eFindings on aging and perspective-taking are mixed. Older adults show declines in lab-based affective tasks (Fernandes et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2019a\u003c/span\u003e; Pratt et al., \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1996a\u003c/span\u003e), yet perform comparably to younger adults when motivated (Zhang et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). They tend to use more positive language in narrative retellings (Sullivan et al., \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) and excel in social inference despite visual/spatial declines (De Lillo and Ferguson, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Perspective-taking deficits in aging have been linked to reduced dorsomedial prefrontal cortex activity (Moran et al., \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe dynamic integration theory (Labouvie-Vief, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2003\u003c/span\u003e) posits that emotional and cognitive complexity peaks in midlife and declines in older adulthood, giving way to emotional prioritization (O'Brien et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Age-related declines in executive function, memory, and processing speed (Hambrick et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Salthouse, \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; West, \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e1996\u003c/span\u003e) may underlie diminished cognitive perspective-taking (Fernandes et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2019b\u003c/span\u003e; Maylor et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Pratt et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1996b\u003c/span\u003e), while moral reasoning shifts from intent to outcome (Margoni et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eSocioemotional Aging and Moral Reasoning\u003c/h2\u003e\u003cp\u003eDespite cognitive declines, older adults often show enhanced emotional regulation and social focus (Isaacowitz et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), extending to moral domains (Margoni et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). They report greater relational satisfaction (Diener et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Lang and Carstensen, \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Litwin, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2001\u003c/span\u003e), demonstrate stronger memory for moral events (Narvaez et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), and favor deontological reasoning linked to moral idealism and affective sensitivity (Arutyunova et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e; McNair et al., \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). This paradox\u0026mdash;cognitive decline alongside social-emotional effectiveness\u0026mdash;may reflect task-context differences (Daley et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Marsiske et al., 1995; Spreng and Turner, \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eFindings remain mixed: older adults rate immoral acts as more wrong but show reduced emotional arousal (Lu and Fung, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Electrophysiology suggests diminished emotional engagement (Fernandes et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2019a\u003c/span\u003e), while neuroimaging shows reduced affective but preserved cognitive empathy circuits in aging (Chen et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2014a\u003c/span\u003e). Moral evaluations involve both cognitive and affective processes across brain regions including the amygdala, insula, anterior mid-cingulate cortex (aMCC), temporal pole, right temporoparietal junction (rTPJ), orbitofrontal cortex (OFC), ventromedial and dorsolateral prefrontal cortex (dlPFC), medial prefrontal cortex (mPFC), and striatum (Buckholtz and Marois, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Moll et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). These systems support moral cognition through arousal, valuation, and social reasoning (Decety et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Pascual et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Older adults tend to favor deontological choices, particularly when the decision feels intuitive, but show no age-related difference when utilitarian responses are also intuitive. This bias is associated with enhanced connectivity between the posterior medial core of the default mode network (pmDN) and the dorsal attention network, alongside reduced segregation of the pmDN from the broader brain network (Huang et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Stronger DMN connectivity supported better memory across ages and reduced utilitarian choices in younger adults. In older adults, similar moral choices involved greater DMN\u0026ndash;Salience Network interaction, suggesting age-related shifts in how DMN supports memory and moral decisions (Daley and Kensinger, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eImplicit and Explicit Moral Judgments Across the Lifespan\u003c/h3\u003e\n\u003cp\u003eExplicit moral judgments are deliberate and reflect rational processing grounded in values and norms, while implicit judgments are automatic and emotionally driven (Cameron et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Greenwald et al., 1998). Aging may alter this balance, with older adults possibly leaning more on affective or habitual responses in moral contexts. Explicit attitudes are commonly assessed via tools like the Justice Sensitivity Inventory (Schmitt et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), and implicit attitudes through the Implicit Association Test (IAT). Given age-related slowing, we applied a z-score transformation to the IAT to account for processing speed differences (Faust et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Hummert et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2002\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eTogether, distinctions between cognitive and affective processing, perspective-taking, and implicit versus explicit judgment styles help illuminate age-related changes in moral reasoning. While behavioral studies highlight decision speed and bias shifts, fMRI enables examination of the neural underpinnings. In this study, we used fMRI with a button-trigger, first-person simulation of moral scenarios to explore these mechanisms across age groups (Chen et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Ruby and Decety, \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2001\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eBased on current understanding of age-related shifts in cognitive and affective processing\u0026mdash;characterized by increased socio-affective motivation and decreased cognitive ability\u0026mdash;we hypothesize the following: (1) First-Person Perspective-Taking Reaction Times: Due to the high socio-affective motivation tied to moral and immoral agency (Zhang et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), older adults will perform comparably to younger adults. (2) Moral Evaluations of Observed Actions: Older adults may exhibit heightened praise/blame evaluations of moral/immoral behaviors, consistent with findings of increased sensitivity to victims\u0026rsquo; unpleasantness during evaluations of harmful actions (Chen et al. (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2014c\u003c/span\u003e)). (3) Implicit and Explicit Moral Attitudes: Implicit attitudes, closely tied to automatic affective inclinations (Isaacowitz et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), may increase with age, whereas explicit attitudes, associated with deliberative cognitive evaluation (Chen et al., 2014b; Spreng and Turner, \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), are expected to remain stable or decline in older adults compared to younger adults. (4) fMRI BOLD Responses: We hypothesize reduced activation in affective brain regions such as the anterior insular cortex (AIC) and anterior mid-cingulate cortex (aMCC) related to pain empathy in older adults, with stable activity in the medial prefrontal cortex and posterior superior temporal sulcus associated with cognitive empathy and perceived agency (Chen et al., 2014b). This shift may indicate more efficient valence processing in older adults, accompanied by increased reliance on cognitive brain subsystems and heightened moral sensitivity due to changes in the cognitive-affective balance.\u003c/p\u003e"},{"header":"MATERIALS AND METHODS","content":"\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e\u003ch2\u003eParticipants\u003c/h2\u003e\u003cp\u003eThe study enrolled two groups of right-handed, ethnic Chinese participants from the community: (1) 31 young adults (15 males, aged 20 to 31 years, mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviation (SD): 23\u0026thinsp;\u0026plusmn;\u0026thinsp;2.2); and (2) 32 older adults (11 males, 65 to 80, 69\u0026thinsp;\u0026plusmn;\u0026thinsp;4.0). One older subject was excluded due to excessive head movement during fMRI scanning, leaving \u003cem\u003eN\u003c/em\u003e\u0026thinsp;=\u0026thinsp;62 for the fMRI data analysis. All participants were screened to ensure that they had no history of neurological or psychiatric disorders, had normal or corrected-normal visual acuity, and were taking no medications at the time of testing. Older participants were further screened for medication use and recent surgical procedures. This study's experimental protocols received ethical approval from the Institutional Review Board of National Yang-Ming University (Protocol Number: YM102033). All participants gave written informed consent for the study, which was approved by local ethic committee, and was conducted in accordance with the \u003cem\u003eDeclaration of Helsinki\u003c/em\u003e.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eVisual Stimuli\u003c/h3\u003e\n\u003cp\u003eForty-five validated animations from previous fMRI studies were presented to participants (Chen et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e; Chen et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020b\u003c/span\u003e; Cheng et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2021a\u003c/span\u003e; Decety and Porges, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Each animation comprised three images with duration of 1000, 200, and 1000 milliseconds, respectively, and which portrayed the following scenarios: (1) a person who is taking an action to physically harming one another person (harming); and (2) a person who is alleviating physical pain from a suffering person (helping). One additional baseline stimuli depicted people carrying out an action that was irrelevant to the other one (neutral).\u003c/p\u003e\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003eProcedures\u003c/h2\u003e\u003cp\u003eParticipants performed moral judgments on moral dilemmas and completed the dispositional Justice Sensitivity Inventory (JSI) and the morally-laden implicit association test (mIAT) before fMRI scanning (supplementary Figure \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003es1\u003c/span\u003e). During fMRI scanning, we made use of mental simulations of social interactions, and a button-trigger design was adopted with reference to the current literature (Chen et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Participants first imagined themselves in the scenario and pressed a handheld button when ready for the action to proceed. They viewed an initial image of the action, then pressed the button to trigger the following two images. There were two runs\u0026mdash;one with simulated harming and the other with helping actions\u0026mdash;intermixed with neutral action blocks. Before each run, participants were instructed to mentally simulate themselves as the agent of either harming or helping actions from a first-person perspective. Participants could press the button at their own pace (mean reaction time\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error: young adults, 1238\u0026thinsp;\u0026plusmn;\u0026thinsp;172 ms; older adults, 1798\u0026thinsp;\u0026plusmn;\u0026thinsp;170 ms), which initiated the complete action sequence. The task aimed to measure the time participants needed to mentally prepare for simulating the actions. Quicker responses suggested easier engagement and lower cognitive load. Before the formal trials, participants practiced with three trials and feedback to ensure understanding. The button-press was designed to help immerse participants in the virtual agent's role. After fMRI scanning, the visual stimuli were presented, and participants, with the use of a computer-based seven-point visual analogue scale, were asked to evaluate how much praise they attributed to themselves as the imagined protagonist in the helping actions or how much blame they attributed to themselves as the imagined protagonist in the harming actions.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eDispositional justice sensitivity (JSI)\u003c/h3\u003e\n\u003cp\u003eThe JSI is a self-reported measure that assesses four aspects of justice sensitivity, with scores ranging from 1 (strongly disagree) to 6 (strongly agree). Three perspectives\u0026mdash;observer, beneficiary, and perpetrator\u0026mdash;are often combined into a single \"other-oriented sensitivity\" score (Edele et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Gollwitzer et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). While the JSI measured participants\u0026rsquo; sensitivity to injustice, unfairness, or immoral scenarios, self-oriented sensitivity referred to situations in which participants themselves were the victims of injustice, making the sensitivity directed toward themselves. In contrast, other-oriented sensitivity referred to sensitivity toward injustice experienced by others (See supplementary materials for detailed methods).\u003c/p\u003e\n\u003ch3\u003eUtilitarian judgments on moral dilemmas\u003c/h3\u003e\n\u003cp\u003eBased on previous work (Chen et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Greene et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Greene et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Greene et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Huebner et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.cell.com/neuron/supplemental/S0896-6273(04)00634-8\u003c/span\u003e\u003cspan address=\"http://www.cell.com/neuron/supplemental/S0896-6273(04)00634-8\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), forty-eight moral dilemmas were selected to create two versions of the moral judgment task balanced for emotional intensity (Koenigs et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Each version included 12 dilemmas: six personal (e.g., pushing a stranger in the trolley dilemma) and six impersonal (e.g., flipping a switch). Personal dilemmas involved direct harm, while impersonal involved indirect harm. Personal dilemmas were further classified as involving inevitable or evitable harm, with higher moral permissibility judgments for inevitable harm. The dilemmas were translated from English to Chinese and back-translated for accuracy. Participants read and responded at their own pace.\u003c/p\u003e\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003eImplicit moral attitudes (mIAT)\u003c/h2\u003e\u003cp\u003eThe mIAT was modified with animations and words for the stimuli. Verbal stimuli included 26 extremely pleasant and 26 extremely unpleasant words, selected from high-frequency Chinese words (Chen et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e; Chen et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Chen et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020b\u003c/span\u003e; Mart\u0026iacute;nez et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The mIAT performance, as indexed by D scores, represents implicit moral attitudes. The D score was calculated by subtracting the mean RT of congruent (immoral-negative) blocks from that of incongruent (immoral-positive) blocks and dividing it by the pooled SD across the two blocks (Nosek et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Higher D scores might come from higher RTs for incongruent blocks or lower RTs for congruent blocks (See supplementary materials for detailed methods).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\u003ch2\u003efMRI acquisition, data processing and analysis\u003c/h2\u003e\u003cp\u003eScanning employed a mixed design (19.1 s ON/13.2\u0026thinsp;\u0026plusmn;\u0026thinsp;4.4 s OFF) with two runs, each consisting of six ON blocks (two each of harming, helping, and neutral scenarios) and six OFF blocks. ON blocks included five trials, each lasting 2200 ms, with pseudo-randomized sequence presentation. Scanning was performed using a 3T Siemens Magnetom Trio-Tim MRI, and BOLD fMRI signals were collected. Functional and structural images were processed using SPM12 and analyzed in MATLAB. Functional images were realigned, co-registered, normalized to MNI space, and smoothed. A voxel-by-voxel regression analysis modeled three conditions (harming, helping, and neutral) to capture neuro-hemodynamic responses, with contrasts for harming vs. neutral and helping vs. neutral. These first-level contrast images were entered into an ANOVA to examine age and scenario effects, with whole-brain activation corrected for multiple comparisons (FWE rate p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). To assess how aging moderated neural responses, ROI data from cognitive and affective networks were extracted based on predefined coordinates from the literature and prior studies. Four cognitive ROIs (dlPFC, dmPFC, TPJ, OFC) and four affective ROIs (amygdala, insula, aMCC, temporal pole) were analyzed using MarsBaR in SPM12 (Chen et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Cheng et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2021b\u003c/span\u003e; Moran et al., \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) (See supplementary materials for detailed methods).\u003c/p\u003e\u003c/div\u003e"},{"header":"RESULTS","content":"\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003ch2\u003eBehavioral Performance\u003c/h2\u003e\u003cdiv id=\"Sec15\" class=\"Section3\"\u003e\u003ch2\u003eDemographics\u003c/h2\u003e\u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e lists demographic data. Although younger participants scored higher on the Mini-Mental Status Examination (MMSE) (young: 29.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1; old: 28.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;4.49, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), none of the older participants were suffering from dementia, as indicated by all MMSE scores being higher than 25 (ranging from 27 to 30) (Folstein et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1975\u003c/span\u003e). No subjects had any evidence of depression, as indicated by scores lower than 5 on the Chinese version of the Geriatric Depression Scale (young: 2.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37; old: 1.47\u0026thinsp;\u0026plusmn;\u0026thinsp;0.25, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.47, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.15) (Mui, \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e1996\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eDemographic and dispositional variables of the participants.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003eYoung\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eOld\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e(\u003cem\u003eN\u003c/em\u003e\u0026thinsp;=\u0026thinsp;31)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003e(\u003cem\u003eN\u003c/em\u003e\u0026thinsp;=\u0026thinsp;32)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMean\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSE\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMean\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eSE\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003ep\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eAge\u003c/b\u003e (years)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e23.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e69.28\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eGender\u003c/b\u003e (male%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e48\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e9.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e34\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.27\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eMMSE\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e29.77\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e28.78\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eDepression\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.37\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.25\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eEducation\u003c/b\u003e (years)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e16.97\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.33\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e14.32\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u0026lt; 0.001\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003emIAT D-score\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.61\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.91\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eJSI\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSelf-oriented\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2.87\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.004\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eOther-oriented\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4.46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.41\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.84\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eJSI total score\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4.27\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.21\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"6\" nameend=\"c6\" namest=\"c1\"\u003e\u003cp\u003e\u003cb\u003eMoral dilemmas\u003c/b\u003e (endorsement %)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNon-moral\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e63.08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5.86\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e56.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4.79\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.39\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eImpersonal\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e52.26\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e7.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e60.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e7.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.45\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eInevitable\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e52.42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e6.49\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e46.88\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e6.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.54\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEvitable\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e20.43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.28\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e25.52\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.17\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"6\" nameend=\"c6\" namest=\"c1\"\u003e\u003cp\u003e\u003cb\u003eMoral evaluation\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBlame\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5.08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.19\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.26\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.006\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePraise\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"6\"\u003eAbbreviations: SE, standard error; MMSE, Mini-Mental Status Examination; mIAT, morality implicit association test; JSI, Justice Sensitivity Inventory; Depression, Chinese version of the Geriatric Depression Scale.\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Sec16\" class=\"Section2\"\u003e\u003ch2\u003eAge differences in implicit moral attitudes\u003c/h2\u003e\u003cp\u003eFor implicit moral attitudes, older adults exhibited higher D scores than young adults (young: 0.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05; old: 0.94\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06, \u003cem\u003et\u003c/em\u003e = -3.59, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA). Because the IAT relies on RTs, higher D scores found in older adults not only reflected implicit attitudes but also age differences in processing speeds. Given that older adults exhibited a greater increase in reaction times (RTs) compared to younger adults, it is essential to disentangle the effect of implicit attitudes from the general age-related slowing. Accordingly, trial-level z-score transformations were applied separately to congruent and incongruent trials (Hummert et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Resulting z-score transformation latencies were then entered into an ANOVA: 2 (age: young vs. old) \u0026times; 2 (congruency: congruent vs. incongruent). Results showed a main effect of congruency (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 61\u003c/sub\u003e = 343.51, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.85; congruent vs. incongruent: -0.38\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 vs. 0.39\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02, mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE) and an interaction effect between age and congruency (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 61\u003c/sub\u003e = 15.46, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.20), while the main effect of age (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 61\u003c/sub\u003e = 1.85, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.18) did not reached significance. Post-hoc tests revealed that, when compared to young adults, older adults showed significant shorter latencies in the congruent trials (young vs. old: -0.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.027 vs. -0.46\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 s, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3.84, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), but longer latencies in the incongruent trials (young vs. old: 0.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.027 vs. 0.48\u0026thinsp;\u0026plusmn;\u0026thinsp;0.032 s, \u003cem\u003et\u003c/em\u003e = -3.99, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec17\" class=\"Section2\"\u003e\u003ch2\u003eAge differences in dispositional justice sensitivity\u003c/h2\u003e\u003cp\u003eThe 2 (age: young vs. old; a between-subject factor) \u0026times; 2 (perspective: self-oriented vs. other-oriented; a within-subject factor) mixed ANOVA on JSI scores showed a main effect of age (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 61\u003c/sub\u003e = 5.25, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.025, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.079; young vs. old: 4.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14 vs. 3.64\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14), perspective (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 61\u003c/sub\u003e = 44.99, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.424; self- vs. other-oriented: 3.28\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14 vs. 4.44\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12), and an interaction effect between age and perspective (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 61\u003c/sub\u003e = 5.3, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.025, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.08). Post-hoc tests revealed that, when compared to young adults, older adults showed significantly reduced victim sensitivity toward self-oriented concerns (young vs. old: 3.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18 vs. 2.87\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2.95, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.004), but comparable sensitivities toward other-oriented concerns (young vs. old: 4.46\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15 vs. 4.41\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.21, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.84) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec18\" class=\"Section2\"\u003e\u003ch2\u003eAge differences in moral evaluations\u003c/h2\u003e\u003cp\u003eDue to the RT data not being normally distributed, the RTs were base-10 log-transformed for further analyses. A 2 (age: young vs. old) \u0026times; 3 (scenario: harming vs. helping vs. neutral) repeated ANOVA on log-transformed RTs revealed a main effect of scenario (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e2, 122\u003c/sub\u003e = 51.11, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.46) with the longest RTs for harming behavior (3.19\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 s) as compared to helping (3.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 s, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;9.52, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and neutral actions (3.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 s, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;8.41, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). There was neither a main effect of age (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 61\u003c/sub\u003e = 1.26, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.27) nor an age \u0026times; scenario interaction (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 61\u003c/sub\u003e = 0.01, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.92) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eSubjective ratings included praise for interpersonal assistance and blame for harming behaviors. Compared to younger participants, older adults attributed more praise when helping others (young vs. old: 4.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22 vs. 6.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18, \u003cem\u003et\u003c/em\u003e = -5.72, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and more blame when harming others (young vs. old: 5.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19 vs. 6.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.26, \u003cem\u003et\u003c/em\u003e = -2.85, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.006) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eB).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec19\" class=\"Section2\"\u003e\u003ch2\u003eAge differences in moral dilemmas\u003c/h2\u003e\u003cp\u003eThe 2 (age: young vs. old; a between-subject factor) \u0026times; 4 (dilemma type: nonmoral, impersonal, personal-evitable, or personal-inevitable; a within-subject factor) mixed ANOVA on the endorsement of utilitarian decisions produced a main effect of dilemma type (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e3, 183\u003c/sub\u003e = 20.33, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.25), while there was neither an effect of age (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 61\u003c/sub\u003e = 0.002, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.97) nor age \u0026times; dilemma type interaction (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e3, 183\u003c/sub\u003e = 0.97, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.36). The evitable harm (22.98\u0026thinsp;\u0026plusmn;\u0026thinsp;1.85) was judged as less morally endorsed than did the dilemmas with nonmoral (59.84\u0026thinsp;\u0026plusmn;\u0026thinsp;3.78, \u003cem\u003et\u003c/em\u003e = -10.19, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), impersonal (56.13\u0026thinsp;\u0026plusmn;\u0026thinsp;5.05, \u003cem\u003et\u003c/em\u003e = -7.2, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and the inevitable harm (49.65\u0026thinsp;\u0026plusmn;\u0026thinsp;4.46, \u003cem\u003et\u003c/em\u003e = -5.75, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec20\" class=\"Section2\"\u003e\u003ch2\u003eNeuroimaging results\u003c/h2\u003e\u003cdiv id=\"Sec21\" class=\"Section3\"\u003e\u003ch2\u003eNeuro-hemodynamic response to moral scenarios\u003c/h2\u003e\u003cp\u003eThe entire sample of 62 participants (regardless of age) showed significant neuro-hemodynamic increases in the network of regions involved in both cognitive and affective processing of moral brain (\u003cem\u003ek\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;20, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05, FWE-corrected) (Healey and Grossman, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Regions with greater activity in response to scenarios involving interpersonal harm vs. assistance included the dmPFC, dlPFC, TPJ, OFC, insula, aMCC, temporal pole, hippocampus, caudate, amygdala, postcentral gyrus, and occipital cortex. The reverse contrast showed no significant cluster (supplementary Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003es1\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Sec22\" class=\"Section2\"\u003e\u003ch2\u003eROI results\u003c/h2\u003e\u003cp\u003eRegarding the age effect on neuro-hemodynamic responses, while there was no survival cluster of the main effect of age (young vs. old) found in the voxel-wise analyses, the interaction between age (young vs. old) and scenario (harming vs. helping) was identified among several ROIs, including the dlPFC (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 60\u003c/sub\u003e = 4.1, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.047, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.06), OFC (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 60\u003c/sub\u003e = 4.77, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.033, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.07), insula (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 60\u003c/sub\u003e = 4.14, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.046, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.07), and aMCC (\u003cem\u003eF\u003c/em\u003e\u003csub\u003e1, 60\u003c/sub\u003e = 5.83, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.019, pη\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.09) (supplementary Table s2).\u003c/p\u003e\u003cp\u003ePost-hoc analyses indicated that scenario effects in the dlPFC, OFC, insula, and aMCC showed opposite directions depending on the factor of age. Harming compared to helping actions showed increased activity in the aMCC (harming vs. helping: 0.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11 vs. -0.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2.85, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.008) with a trend in the insula (0.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09 vs. -0.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.75, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.09) in young adults, but none in older adults (insula: -0.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 vs. 0.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07, \u003cem\u003et\u003c/em\u003e = -1.04, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.306; aMCC: 0.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08 vs. 0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07, \u003cem\u003et\u003c/em\u003e = -0.06, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.955). On the other hand, harming as compared to helping actions showed increased activities in the dlPFC (0.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12 vs. -0.15\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2.16, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.039) and OFC (0.38\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19 vs. -0.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19, \u003cem\u003et\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2.43, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.021) in older adults, but none in young adults (dlPFC: -0.002\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13 vs. 0.15\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1, \u003cem\u003et\u003c/em\u003e = -0.82, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.418; OFC: 0.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2 vs. 0.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18, \u003cem\u003et\u003c/em\u003e = -0.89, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.38) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eRegarding brain-behavior correlations, the young and older groups showed a double dissociation between neural correlates and behavioral assessments (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). When adopting harming actions, the amygdala activity positively predicted D scores of implicit moral attitudes in young adults (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.43, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.015), but not in older adults (\u003cem\u003er\u003c/em\u003e = -0.25, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.18). The rTPJ activity negatively predicted the self-related justice sensitivity within the young (r= -0.502, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.004), but not within the elders (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.29, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.11). The temporal pole activity positively predicted the blame ratings in young adults (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.8, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.038), but not in older adults (\u003cem\u003er\u003c/em\u003e = -0.03, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.87). Fisher\u0026rsquo;s z tests confirmed the significant dissociations in the amygdala (z\u0026thinsp;=\u0026thinsp;2.68, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.007) and rTPJ (z\u0026thinsp;=\u0026thinsp;3.18, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.002). On the other hand, adopting helping actions had no such dissociation.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"DISCUSSION","content":"\u003cp\u003eThis study revealed age-related changes in implicit and explicit moral attitudes and their neural correlates during perspective-taking of moral behavior. While aging had no effect on moral dilemmas, older adults exhibited higher D scores on the mIAT and lower victim sensitivity on the JSI. Reaction times for social perspective-taking showed no age-related differences, but older adults attributed greater praise and blame to moral behaviors. Reduced differential BOLD activity was observed in the insula and anterior mid-cingulate cortex (aMCC) for helping versus harming, while the orbitofrontal cortex (OFC) and dorsolateral prefrontal cortex (dlPFC) maintained differentiation. Unlike younger adults, correlations between BOLD activity and moral attitudes were absent in older adults. These findings suggest greater moral cognitive evaluation in older adults, reflecting a neural shift in cognitive and affective processing with aging.\u003c/p\u003e\u003cp\u003eContrary to expectations, moral dilemmas showed no age effects or interactions, potentially due to cultural differences. Increased deontological judgments in older adults are more prominent in Western cultures (Arutyunova et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e). Chinese individuals are less likely to sacrifice one to save five (Gold et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), judging avoidable harm more harshly than impersonal or inevitable harm. Cultural differences between East and West influence moral concepts. In collectivist Chinese culture, morality emphasizes social harmony and respect for hierarchy, extending to behaviors deemed impolite in Western contexts, such as not offering a seat (Nichols, 2022). By contrast, Western individualistic cultures prioritize utilitarian ethics, favoring decisions maximizing collective welfare, even with trade-offs (Arutyunova et al., 2016b; Bentahila et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Hong and Sciences, 2023). These cultural norms may explain why age-related shifts in moral reasoning observed in Western contexts are less pronounced in Chinese culture, where Confucian values persist across age groups.\u003c/p\u003e\u003cp\u003eOlder adults showed significantly higher D scores on the implicit moral attitudes, as indicated by the mIAT, even after controlling for age-related slowing effects on RTs. The higher D scores might have resulted from either higher RTs for incongruent (immoral-positive) blocks or lower RTs for congruent (immoral-negative) blocks (Nosek et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). The D scores on the mIAT were predictive of actor-observer bias when attributing blame for immoral behaviors and individual differences during moral computation (Chen et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e; Chen et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Chen et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020b\u003c/span\u003e). Higher D scores, revealing increased conflict between positive/negative valence and immoral/moral behaviors, might indicate that older adults experienced more difficulty in distinguishing the greater cognitive evaluation triggered by moral behaviors from the conflict arising from word valence pairs. In parallel, they also explicitly attributed more praise and blame to their own moral behaviors. Additionally, older adults reported lower victim sensitivity toward self-oriented concerns but comparable sensitivity toward other-oriented concerns on the JSI. This is supported by previous findings indicating that older adults tend to exhibit more morally idealistic beliefs (Margoni et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). These behavioral results support our hypotheses: (1) Older adults, driven by high socio-affective motivation related to moral and immoral agency, demonstrated performance comparable to younger adults, with no significant differences observed in reaction times. (2) Consistent with previous findings, older adults exhibited increased sensitivity, assigning heightened praise or blame to moral and immoral behaviors relative to younger adults. (3) Older adults showed stronger implicit moral attitudes, indicating greater conflict between automatic affective inclinations toward positive or negative valence and moral or immoral actions. Interestingly, on explicit measures of justice sensitivity, older adults showed reduced sensitivity when they themselves were the victims of injustice, reflecting diminished self-oriented sensitivity, while maintaining sensitivity toward injustices experienced by others. This pattern may be partially attributed to high socio-affective motivation and the cultural framework discussed above.\u003c/p\u003e\u003cp\u003eIn contrast to young adults, older adults retained differential BOLD activities in the OFC and dlPFC in response to harming vs. helping others, but exhibited reduced differential activities in the insula and aMCC. When perceiving moral scenarios, older relative to younger adults showed an intent-to-outcome shift specific to harming actions, which was associated with a decline in theory of mind abilities (Margoni et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In line with age-related changes in empathy for pain, activity in the anterior insula and anterior mid-cingulate cortex (aMCC) exhibited age-related declines, whereas activity in the medial prefrontal cortex (mPFC) and right temporoparietal junction (rTPJ) remained preserved in older adults, even as they reported increased unpleasantness (Chen et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2014c\u003c/span\u003e). Considering the heightened behavioral sensitivity and reduced activity in the anterior insular cortex (AIC) and anterior mid-cingulate cortex (aMCC) associated with the affective components of pain empathy and affective perspective-taking in older adults, these findings\u0026mdash;alongside increased activity observed in the orbitofrontal cortex (OFC) and right temporoparietal junction (rTPJ)\u0026mdash;may indicate a shifting balance between cognitive and affective processing with age (Healey and Grossman, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). This shift suggests that valence processing within the affective brain system may become more efficient in older adults, leading to reduced activation in these regions. Consequently, older adults may allocate more resources to cognitive brain subsystems, which typically show age-related declines (Brehmer et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Kensinger et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eCorrelations between BOLD activities and moral attitudes and ratings found in young adults did not exist in older adults. Based on the same paradigm, young adults revealed that the OFC mediated the mIAT\u0026ndash;RTs association, and the rTPJ mediated the JSI\u0026ndash;warm-glow linkage (Chen et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Young adults also displayed positive correlations of the mIAT with neural responses in the OFC and temporal poles to actor-observer asymmetry (Chen et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e). Such brain-behavior correlations vanished with aging. One previous fMRI study revealed that older adults frequently exhibited greater magnitudes and extents of activation of the BOLD signals compared to younger adults. This additional activation might reflect compensatory recruitment associated with functional and structural deterioration of neural resources (Woodard and Sugarman, \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn interpreting our findings, it is essential to consider the nature of the moral task used in this study. While participants were instructed to mentally simulate themselves as agents in morally laden actions, the button-trigger design with sequential image presentation places the task at an intersection between first-person and third-person moral judgment paradigms. Prior work has shown that first-person tasks tend to elicit greater emotional engagement and activation in affective networks (Greene et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), whereas third-person tasks are more likely to involve reflective perspective-taking and recruit cognitive control regions (Koenigs et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Our paradigm, with its explicit instruction for self-agency but lack of real-world consequence, may have evoked a hybrid engagement, consistent with findings that moral cognition involves both intuitive and deliberative processes (Daley and Kensinger, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Huang et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Age-related neural differences observed here\u0026mdash;specifically, the diminished insula and aMCC response in older adults alongside preserved OFC and dlPFC activity\u0026mdash;may reflect a shift in the balance of these networks, supporting the idea that aging modulates the integration of affective and cognitive moral systems(Daley and Kensinger, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). These distinctions underscore the importance of carefully considering task framing in moral neuroscience research, particularly when comparing across age groups.\u003c/p\u003e\u003cp\u003eA few limitations of the current work should be clarified for future research. For instance, we adopted a first-person perspective, rather than a third-person perspective, thereby limiting direct comparisons with previous available literatures. First- or third-person perspective in moral cognition involves distinct neural processes (Avram et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Chen et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e). In young adults, dlPFC and rTPJ activities were involved in the asymmetry between first-hand experiencing and passive viewing moral behaviors, and the first-person relative to third-person perspective elicited stronger activity in the aMCC (Chen et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e). To discern any aging impact on the neural underpinnings of moral behavior, it would be better to use a first-person perspective. Finally, our sample included only young and older adults, but did not include middle-aged adults. This might not be an optimal design, and future studies are warranted with longitudinal research in clinical populations or large community-based samples.\u003c/p\u003e\u003cp\u003eAll in all, this study demonstrated that aging has an impact on the neural underpinnings of moral attitudes and behaviors. The results suggest a reduced engagement of neural affective subsystems, with older adults relying more on the neural cognitive subsystems of the prefrontal cortex. These findings could contribute to the existing body of developmental research (Cowell and Decety, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Decety et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) in demonstrating the neuro-cognitive trajectories of morality over a person\u0026rsquo;s lifespan. Given the fact that morality arguably serves adaptive social functions during one\u0026rsquo;s lifespan, especially in adulthood as individuals become responsible for their own and for others well-being (Gebremariam and Sadana, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), the present findings may contribute to a better understanding of healthy aging, and help provide insights into neurodegenerative disorders with socio-emotional deficits.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eACKNOWLEDGMENTS\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank Jwu-Jiun Chang for assisting with data collection and preparation of stimuli. We thank Prof. Jean Decety for the morally-laden scenarios used as the stimuli; without his generous contribution such an endeavor would not have been possible.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFUNDINGS\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe study was funded by the National Science and Technology Council, R.O.C. (NSTC 113-2410-HA49-065; 113-2410-H-038-034 -; 113-2410-H-028 -005 -), National Yang Ming Chiao Tung University Hospital (RD2025-002), and the Higher Education Sprout Project by the Ministry of Education (MOE) in Taiwan (TMU-114-N-05), Taiwan.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCONFLICT OF INTEREST\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNone of the authors has any conflicts of interest to declare.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDATA AVAILABILITY STATEMENT\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe data presented in this study are available on request from the corresponding author.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCRediT authorship contribution statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eC.C. and Y.C. conceived and designed the study. C.C. and Y.C. supervised the study. C.C. and Y.C. wrote the main manuscript text. Y.C.C. curated the data. C.C. formally analyzed the data. C.C., Y.C.C., Y.T.F., R.M.M., and Y.C. reviewed and edited the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of Generative AI and AI-Assisted Technologies:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe author(s) acknowledge the use of generative AI and AI-assisted technologies during the preparation of this manuscript. Specifically, OpenAI’s ChatGPT was used to improve the clarity, grammar, and readability of the English language in the writing process. No content was generated or interpreted by the AI; all conceptual, analytical, and interpretative work was conducted by the authors. The authors take full responsibility for the integrity and accuracy of the content presented.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eArutyunova, K.R., Alexandrov, Y.I., Hauser, M.D., 2016a. Sociocultural Influences on Moral Judgments: East-West, Male-Female, and Young-Old. Front Psychol 7, 1334.\u003c/li\u003e\n\u003cli\u003eArutyunova, K.R., Alexandrov, Y.I., Hauser, M.D.J.F.i.p., 2016b. Sociocultural influences on moral judgments: East\u0026ndash;west, male\u0026ndash;female, and young\u0026ndash;old. 7, 1334.\u003c/li\u003e\n\u003cli\u003eAvram, M., Hennig-Fast, K., Bao, Y., Poppel, E., Reiser, M., Blautzik, J., Giordano, J., Gutyrchik, E., 2014. Neural correlates of moral judgments in first- and third-person perspectives: implications for neuroethics and beyond. BMC Neurosci 15, 39.\u003c/li\u003e\n\u003cli\u003eBentahila, L., Fontaine, R., Pennequin, V.J.F.i.P., 2021. Universality and cultural diversity in moral reasoning and judgment. 12, 764360.\u003c/li\u003e\n\u003cli\u003eBrehmer, Y., Westerberg, H., B\u0026auml;ckman, L.J.F.i.h.n., 2012. Working-memory training in younger and older adults: training gains, transfer, and maintenance. 6, 63.\u003c/li\u003e\n\u003cli\u003eBuckholtz, J.W., Marois, R., 2012. The roots of modern justice: cognitive and neural foundations of social norms and their enforcement. Nat Neurosci 15, 655-661.\u003c/li\u003e\n\u003cli\u003eCameron, C.D., Payne, B.K., Sinnott-Armstrong, W., Scheffer, J.A., Inzlicht, M., 2017. Implicit moral evaluations: A multinomial modeling approach. Cognition 158, 224-241.\u003c/li\u003e\n\u003cli\u003eChen, C., Decety, J., Huang, P.C., Chen, C.Y., Cheng, Y., 2016. Testosterone administration in females modulates moral judgment and patterns of brain activation and functional connectivity. Human Brain Mapping 37, 3417-3430.\u003c/li\u003e\n\u003cli\u003eChen, C., Martinez, R.M., Chen, Y., Cheng, Y., 2020a. Pointing fingers at others: The neural correlates of actor-observer asymmetry in blame attribution. Neuropsychologia 136, 107281.\u003c/li\u003e\n\u003cli\u003eChen, C., Martinez, R.M., Chen, Y.C., Fan, Y.T., Cheng, Y., 2022. The neural mediators of moral attitudes and behaviors. Behav Brain Res 430, 113934.\u003c/li\u003e\n\u003cli\u003eChen, C., Martinez, R.M., Cheng, Y., 2020b. The key to group fitness: The presence of another synchronizes moral attitudes and neural responses during moral decision-making. Neuroimage 213, 116732.\u003c/li\u003e\n\u003cli\u003eChen, Y.-C., Chen, C.-C., Decety, J., Cheng, Y., 2014a. Aging is associated with changes in the neural circuits underlying empathy. Neurobiology of aging 35, 827-836.\u003c/li\u003e\n\u003cli\u003eChen, Y.-C., Chen, C.-C., Decety, J., Cheng, Y.J.N.o.a., 2014b. Aging is associated with changes in the neural circuits underlying empathy. 35, 827-836.\u003c/li\u003e\n\u003cli\u003eChen, Y.C., Chen, C.C., Decety, J., Cheng, Y., 2014c. Aging is associated with changes in the neural circuits underlying empathy. Neurobiol Aging 35, 827-836.\u003c/li\u003e\n\u003cli\u003eCheng, Y., Chou, J., Martinez, R.M., Fan, Y.T., Chen, C., 2021a. Psychopathic traits mediate guilt-related anterior midcingulate activity under authority pressure. Sci Rep 11, 14856.\u003c/li\u003e\n\u003cli\u003eCheng, Y., Chou, J., Mart\u0026iacute;nez, R.M., Fan, Y.-T., Chen, C., 2021b. Psychopathic traits mediate guilt-related anterior midcingulate activity under authority pressure. Scientific reports 11, 1-11.\u003c/li\u003e\n\u003cli\u003eCowell, J.M., Decety, J., 2015. The neuroscience of implicit moral evaluation and its relation to generosity in early childhood. Curr Biol 25, 93-97.\u003c/li\u003e\n\u003cli\u003eDaley, R.T., Kensinger, E.A.J.A., Neuropsychology,, Cognition, 2023. Cognitive decline, socioemotional change, or both? How the science of aging can inform future research on sacrificial moral dilemmas. 30, 272-299.\u003c/li\u003e\n\u003cli\u003eDaley, R.T., Kensinger, E.A.J.N., 2022. Age-related differences in Default Mode Network resting-state functional connectivity but not gray matter volume relate to sacrificial moral decision-making and working memory performance. 177, 108399.\u003c/li\u003e\n\u003cli\u003eDe Lillo, M., Ferguson, H.J.J.J.o.E.P.G., 2023. Perspective-taking and social inferences in adolescents, young adults, and older adults. 152, 1420.\u003c/li\u003e\n\u003cli\u003eDecety, J., Michalska, K.J., Kinzler, K.D., 2012. The contribution of emotion and cognition to moral sensitivity: a neurodevelopmental study. Cereb Cortex 22, 209-220.\u003c/li\u003e\n\u003cli\u003eDecety, J., Porges, E.C., 2011. Imagining being the agent of actions that carry different moral consequences: an fMRI study. Neuropsychologia 49, 2994-3001.\u003c/li\u003e\n\u003cli\u003eDiener, E., Suh, E.M., Lucas, R.E., Smith, H.L., 1999. Subjective well-being: Three decades of progress. Psychological Bulletin 125, 276-302.\u003c/li\u003e\n\u003cli\u003eEdele, A., Dziobek, I., Keller, M., 2013. Explaining altruistic sharing in the dictator game: the role of affective empathy, cognitive empathy, and justice sensitivity. Learning and Individual Differences 24.\u003c/li\u003e\n\u003cli\u003eFaust, M.E., Balota, D.A., Spieler, D.H., Ferraro, F.R.J.P.b., 1999. Individual differences in information-processing rate and amount: implications for group differences in response latency. 125, 777.\u003c/li\u003e\n\u003cli\u003eFernandes, C., Gon\u0026ccedil;alves, A., Pasion, R., Ferreira-Santos, F., Barbosa, F., Martins, I., Marques-Teixeira, J., 2019a. Age-related decline in emotional perspective-taking: Its effect on the late positive potential. Cognitive, Affective, Behavioral Neuroscience 19, 109-122.\u003c/li\u003e\n\u003cli\u003eFernandes, C., Goncalves, A.R., Pasion, R., Ferreira-Santos, F., Barbosa, F., Martins, I.P., Marques-Teixeira, J., 2019b. Age-related decline in emotional perspective-taking: Its effect on the late positive potential. Cogn Affect Behav Neurosci 19, 109-122.\u003c/li\u003e\n\u003cli\u003eFolstein, M.F., Folstein, S.E., McHugh, P.R., 1975. \u0026quot;Mini-mental state\u0026quot;. A practical method for grading the cognitive state of patients for the clinician. J Psychiatr Res 12, 189-198.\u003c/li\u003e\n\u003cli\u003eGebremariam, K.M., Sadana, R., 2019. On the ethics of healthy ageing: setting impermissible trade-offs relating to the health and well-being of older adults on the path to universal health coverage. Int J Equity Health 18, 140.\u003c/li\u003e\n\u003cli\u003eGold, N., Colman, A.M., Pulford, B.D., 2014. Cultural differences in responses to real-life and hypothetical trolley problems. Judgment and Decision Making 9, 65-76.\u003c/li\u003e\n\u003cli\u003eGollwitzer, M., Rothmund, T., Pfeiffer, A., Ensenbach, C., 2009. Why and when justice sensitivty leads to pro- and antisocial behavior. Journal of Research in Personality 43, 999-1005.\u003c/li\u003e\n\u003cli\u003eGreene, J.D., Cushman, F.A., Stewart, L.E., Lowenberg, K., Nystrom, L.E., Cohen, J.D., 2009. Pushing moral buttons: The interaction between personal force and intention in moral judgment. Cognition 111, 364-371.\u003c/li\u003e\n\u003cli\u003eGreene, J.D., Nystrom, L.E., Engell, A.D., Darley, J.M., Cohen, J.D., 2004. The neural bases of cognitive conflict and control in moral judgment. Neuron 44, 389-400.\u003c/li\u003e\n\u003cli\u003eGreene, J.D., Sommerville, R.B., Nystrom, L.E., Darley, J.M., Cohen, J.D., 2001. An fMRI investigation of emotional engagement in moral judgment. Science 293, 2105-2108.\u003c/li\u003e\n\u003cli\u003eGreenwald, A.G., McGhee, D.E., Schwartz, J.L.J.J.o.p., psychology, s., 1998. Measuring individual differences in implicit cognition: the implicit association test. 74, 1464.\u003c/li\u003e\n\u003cli\u003eHambrick, D.Z., Salthouse, T.A., Meinz, E.J., 1999. Predictors of crossword puzzle proficiency and moderators of age-cognition relations. J Exp Psychol Gen 128, 131-164.\u003c/li\u003e\n\u003cli\u003eHealey, M.L., Grossman, M., 2018. Cognitive and affective perspective-taking: evidence for shared and dissociable anatomical substrates. Frontiers in neurology 9, 491.\u003c/li\u003e\n\u003cli\u003eHong, H.J.J.o.E., Humanities, Sciences, S., 2023. Cultural Differences in Moral Judgement. 10, 146-151.\u003c/li\u003e\n\u003cli\u003eHuang, S., Faul, L., Sevinc, G., Mwilambwe-Tshilobo, L., Setton, R., Lockrow, A.W., Ebner, N.C., Turner, G.R., Spreng, R.N., De Brigard, F.J.P., aging, 2021. Age differences in intuitive moral decision-making: Associations with inter-network neural connectivity. 36, 902.\u003c/li\u003e\n\u003cli\u003eHuebner, B., Hauser, M.D., Pettit, P., 2011. How the source, inevitability and means of bringing about harm interact in folk‐moral judgments. Mind Language 26, 210-233.\u003c/li\u003e\n\u003cli\u003eHummert, M.L., Garstka, T.A., O\u0026apos;Brien, L.T., Greenwald, A.G., Mellott, D.S., 2002. Using the implicit association test to measure age differences in implicit social cognitions. Psychol Aging 17, 482-495.\u003c/li\u003e\n\u003cli\u003eIsaacowitz, D.M., Freund, A.M., Mayr, U., Rothermund, K., Tobler, P.N.J.T.J.o.G.S.B., 2021. Age-related changes in the role of social motivation: Implications for healthy aging. 76, S115-S124.\u003c/li\u003e\n\u003cli\u003eKensinger, E.A., Gutchess, A.H.J.J.o.G.S.B.P.S., Sciences, S., 2017. Cognitive aging in a social and affective context: Advances over the past 50 years. 72, 61-70.\u003c/li\u003e\n\u003cli\u003eKoenigs, M., Young, L., Adolphs, R., Tranel, D., Cushman, F., Hauser, M., Damasio, A., 2007. Damage to the prefrontal cortex increases utilitarian moral judgements. Nature 446, 908-911.\u003c/li\u003e\n\u003cli\u003eLabouvie-Vief, G., 2003. Dynamic Integration: Affect, Cognition, and the Self in Adulthood. Current Directions in Psychological Science 12, 201-206.\u003c/li\u003e\n\u003cli\u003eLang, F.R., Carstensen, L.L., 1994. Close emotional relationships in late life: further support for proactive aging in the social domain. Psychol Aging 9, 315-324.\u003c/li\u003e\n\u003cli\u003eLitwin, H., 2001. Social network type and morale in old age. Gerontologist 41, 516-524.\u003c/li\u003e\n\u003cli\u003eLu, M., Fung, H.H., 2019. Moral gain or decay? examing age-related changes in moral judgment. Innov Aging 3, S785.\u003c/li\u003e\n\u003cli\u003eMargoni, F., Geipel, J., Hadjichristidis, C., Bakiaj, R., Surian, L.J.C.s., 2023. Age‐Related Differences in Moral Judgment: The Role of Probability Judgments. 47, e13345.\u003c/li\u003e\n\u003cli\u003eMargoni, F., Geipel, J., Hadjichristidis, C., Surian, L., 2018. Moral Judgment in Old Age. Exp Psychol 65, 105-114.\u003c/li\u003e\n\u003cli\u003eMarsiske, M., Willis, S.L.J.P., aging, 1995. Dimensionality of everyday problem solving in older adults. 10, 269.\u003c/li\u003e\n\u003cli\u003eMart\u0026iacute;nez, R.M., Chen, C.-Y., Liao, T.-T., Cheng, Y., Fan, Y.-T., Chou, S.-H., Chen, C.J.F.i.P., 2020. The multifaceted effects of serotonin transporter polymorphism (5-HTTLPR) on anxiety, implicit moral attitudes, and harmful behaviors. 11, 1521.\u003c/li\u003e\n\u003cli\u003eMaylor, E.A., Moulson, J.M., Muncer, A.M., Taylor, L.A., 2002. Does performance on theory of mind tasks decline in old age? Br J Psychol 93, 465-485.\u003c/li\u003e\n\u003cli\u003eMcNair, S., Okan, Y., Hadjichristidis, C., de Bruin, W.B., 2018. Age differences in moral judgment: older adults are more deontological than younger adults. Behavioral Decsion Making 32, 47-60.\u003c/li\u003e\n\u003cli\u003eMoll, J., de Oliveira-Souza, R., Bramati, I.E., Grafman, J., 2002. Functional networks in emotional moral and nonmoral social judgments. Neuroimage 16, 696-703.\u003c/li\u003e\n\u003cli\u003eMoran, J.M., Jolly, E., Mitchell, J.P., 2012. Social-cognitive deficits in normal aging. J Neurosci 32, 5553-5561.\u003c/li\u003e\n\u003cli\u003eMui, A.C., 1996. Geriatric Depression Scale as a community screening instrument for elderly Chinese immigrants. Int Psychogeriatr 8, 445-458.\u003c/li\u003e\n\u003cli\u003eNarvaez, D., Radvansky, G.A., Lynchard, N.A., Copeland, D.E., 2011. Are older adults more attuned to morally charged information? Exp Aging Res 37, 398-434.\u003c/li\u003e\n\u003cli\u003eNosek, B.A., Bar-Anan, Y., Sriram, N., Axt, J., Greenwald, A.G., 2014. Understanding and using the brief Implicit Association Test: recommended scoring procedures. PLoS One 9, e110938.\u003c/li\u003e\n\u003cli\u003eO\u0026apos;Brien, E., Konrath, S.H., Gr\u0026uuml;hn, D., Hagen, A.L., 2013. Empathic concern and perspective taking: linear and quadratic effects of age across the adult life span. J Gerontol B Psychol Sci Soc Sci 68, 168-175.\u003c/li\u003e\n\u003cli\u003ePascual, L., Rodrigues, P., Gallardo-Pujol, D., 2013. How does morality work in the brain? A functional and structural perspective of moral behavior. Front Integr Neurosci 7, 65.\u003c/li\u003e\n\u003cli\u003ePratt, M.W., Diessner, R., Pratt, A., Hunsberger, B., Pancer, S.M., 1996a. Moral and social reasoning and perspective taking in later life: a longitudinal study. Psychology Aging 11, 66.\u003c/li\u003e\n\u003cli\u003ePratt, M.W., Diessner, R., Pratt, A., Hunsberger, B., Pancer, S.M., 1996b. Moral and social reasoning and perspective taking in later life: a longitudinal study. Psychol Aging 11, 66-73.\u003c/li\u003e\n\u003cli\u003eRuby, P., Decety, J., 2001. Effect of subjective perspective taking during simulation of action: a PET investigation of agency. Nat Neurosci 4, 546-550.\u003c/li\u003e\n\u003cli\u003eSalthouse, T.A., 1994. Aging associations: influence of speed on adult age differences in associative learning. J Exp Psychol Learn Mem Cogn 20, 1486-1503.\u003c/li\u003e\n\u003cli\u003eSchmitt, M., Baumert, A., Gollwitzer, M., Maes, J., 2010. The Justice Sensitivity Inventory: Factorial validity, location in the personality facet space, demorgraphic pattern, and normative data. Social Justice Research 23, 211-238.\u003c/li\u003e\n\u003cli\u003eSpreng, R.N., Turner, G.R.J.P.o.P.S., 2019. The shifting architecture of cognition and brain function in older adulthood. 14, 523-542.\u003c/li\u003e\n\u003cli\u003eSullivan, S.J., Mikels, J.A., Carstensen, L.L.J.P., aging, 2010. You never lose the ages you\u0026rsquo;ve been: Affective perspective taking in older adults. 25, 229.\u003c/li\u003e\n\u003cli\u003eWest, R.L., 1996. An application of prefrontal cortex function theory to cognitive aging. Psychol Bull 120, 272-292.\u003c/li\u003e\n\u003cli\u003eWoodard, J.L., Sugarman, M.A., 2012. Functional Magnetic Resonance Imaging in Aging and Dementia: Detection of Age-Related Cognitive Changes and Prediction of Cognitive Decline, in: Pardon, M.-C., Bondi, M.W. (Eds.), Behavioral Neurobiology of Aging. Springer Berlin Heidelberg, Berlin, Heidelberg, pp. 113-136.\u003c/li\u003e\n\u003cli\u003eZhang, X., Fung, H.H., Stanley, J.T., Isaacowitz, D.M., Ho, M.Y.J.D.P., 2013. Perspective taking in older age revisited: a motivational perspective. 49, 1848.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"aging, moral behavior, moral attitudes, social perspective-taking, fMRI","lastPublishedDoi":"10.21203/rs.3.rs-6859135/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6859135/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eMorality involves both cognitive and affective abilities, and while aging may hamper cognitive perspective-taking, it often enhances social-emotional focus. With strong motivation, older adults can perform as well as younger individuals in affective tasks. However, how aging affects the neural basis of Social Perspective Taking across these systems remains unclear. This study included young and older adults with varying implicit moral attitudes (mIAT) and Justice Sensitivity Inventory (JSI). fMRI scans were conducted while participants mentally simulated moral behaviors, such as helping or harming others. Older adults displayed higher mIAT D scores, reflecting greater conflict between positive valence and immoral actions. They also showed lower victim sensitivity on the JSI and attributed more praise and blame to moral actions, suggesting increased cognitive engagement in evaluating moral behaviors. Neuroimaging revealed that older adults showed reduced neural variation between helping and harming behaviors in the insula and anterior mid-cingulate cortex, indicating less engagement of affective systems. However, they maintained variation in the orbitofrontal and dorsolateral prefrontal cortex, relying more on cognitive systems. These findings suggest older adults compensate for diminished affective responses by engaging cognitive resources during moral evaluations and social perspective-taking, reflecting greater reliance on cognitive processing in differentiating moral behaviors.\u003c/p\u003e","manuscriptTitle":"Aging Changes the Neural Correlates of Social Perspective Taking and Moral Judgements","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-03 01:09:50","doi":"10.21203/rs.3.rs-6859135/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"0292ed09-5fba-4326-aeb0-f02e9cdcaf5d","owner":[],"postedDate":"October 3rd, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":55409959,"name":"Biological sciences/Neuroscience/Cognitive neuroscience"},{"id":55409960,"name":"Biological sciences/Neuroscience/Emotion"},{"id":55409961,"name":"Biological sciences/Neuroscience/Social neuroscience"},{"id":55409962,"name":"Biological sciences/Psychology"},{"id":55409963,"name":"Biological sciences/Psychology/Human behaviour"}],"tags":[],"updatedAt":"2025-11-07T15:38:39+00:00","versionOfRecord":[],"versionCreatedAt":"2025-10-03 01:09:50","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6859135","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6859135","identity":"rs-6859135","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}