Long-term impact of war stress in civilians from the former Yugoslavia: A two-generation neurostructural and psychological study

Long-term impact of war stress in civilians from the former Yugoslavia: A two-generation neurostructural and psychological study | 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 Long-term impact of war stress in civilians from the former Yugoslavia: A two-generation neurostructural and psychological study Monika Fňašková, Pavel Říha, David Ulčák, Marek Preiss, Markéta Nečasová, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7028508/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 17 Mar, 2026 Read the published version in Scientific Reports → Version 1 posted 10 You are reading this latest preprint version Abstract The long-term neurobiological and psychological effects of war-related stress on civilians remain understudied. This study focuses on survivors of the war in the former Yugoslavia (G1) who now reside in the Czech Republic and on the children of survivors (G2) who were born after the conflict. Participants from G1, G2, and a control group (CG) with no war experience underwent structural MRI, answered a semi-structured interview, and completed psychological questionnaires (PTGI, PCL-5, SWSL, MSPSS, Brief-COPE). Voxel-based morphometry was used to assess brain volume differences. Compared to CG, G1 showed reduced grey matter volume in regions associated with PTSD and autobiographical memory, including Crus II, parahippocampal gyrus, middle temporal gyrus, and fusiform gyrus. Psychologically, G1 reported higher PTSD symptoms, lower life satisfaction, and greater post-traumatic growth. G2 showed no structural brain changes but scored higher on post-traumatic growth than CG, with no significant differences in other psychological measures. The findings suggest long-lasting neuroanatomical and psychological effects of war stress in directly exposed individuals (G1). Although G2 showed no brain alterations, the increased post-traumatic growth may indicate subtle adaptation effects of growing up in a post-war environment, but not necessarily only in a maladaptive sense. Biological sciences/Neuroscience Biological sciences/Psychology Social science/Psychology MRI stress civilians war Crus II posttraumatic growth Figures Figure 1 Introduction The conflict in the former Yugoslavia, which lasted from the 1990s to the beginning of the new millennium, was one of the worst and most extensive in Europe since World War II. The wars that successively affected Slovenia, Croatia, Bosnia and Herzegovina, and Kosovo beginning in 1991, and the NATO bombing of Serbia and Montenegro in 1999 in response to the conflict in Kosovo, left the population affected by many traumatic experiences including war injuries, death of loved ones, separation of families, involuntary displacement from their homes, sexual violence, and torture 1 , 2 . Post-war development in these countries devastated by war or bombing has been difficult. Psychological help has not been routinely available, and untreated trauma has often manifested itself in alcohol abuse or violence 3 . The impact of poverty and harsh conditions, and the daily narrative of the war, has taken a toll on the next generation 4 , 5 . This study investigates the long-term effects of war-related stress on civilian populations (G1) and their descendants (G2) from the countries of the former Yugoslavia. Utilising structural magnetic resonance imaging, a psychological survey, and interviews, we explore whether and how the impacts of war have influenced their current lives. Neuroimaging studies investigating the effects of traumatic psychological stress have consistently demonstrated alterations in brain structure and function, including reduced grey matter volume, often associated with the development of PTSD 6 – 9 . Although the hippocampus, amygdala, and prefrontal cortex are commonly identified as the primary stress-sensitive structures, the results vary and many other regions undergo structural or functional changes due to stress 10 , 11 . In addition to the timing, intensity, and duration of psychological stress, one of the main factors affecting the brain appears to be the nature of the traumatic experience 12 . Studies dealing with the effect of war trauma on the alteration of brain structures in non-military personnel are scarce; most studies have been conducted on soldiers or war veterans 13 , 14 . Even though research on civilians is more demanding due to more diverse war experiences, non-existent or inadequate prior medical records, financial demands of research or non-existent research facilities in a war-torn country, and many other reasons, it is an important topic that should be given adequate attention, because unexplored and untreated trauma may have far-reaching effects on individuals and entire societies. In the context of the war in the former Yugoslavia, we found only one MRI paper focused on brain structure volumes conducted on a group of Croatian war veterans who were not professional soldiers. The authors found a significant reduction in right hippocampal volume and a non-significant reduction in left hippocampal volume in this group 15 . The psychological consequences of war include depressive and other mood disorders, anxiety, paranoid ideations, and post-traumatic stress disorder (PTSD) 16 – 18 . PTSD is generally considered to be an illness caused by a traumatic experience. It has been reported that in the general population, PTSD arises as a result of trauma in about 5–10% of cases; in the case of war stress, the incidence of PTSD roughly doubles, and as many as 26% of persons who have remained in a war-afflicted region have been found to have the condition 19 . The condition and its symptomatology are frequently examined in relation to experiences of armed conflict, with documented impacts on both military personnel and civilian populations. In contexts of sustained or repeated exposure to stress characteristic of war and displacement, a more pervasive psychological response may develop, clinically referred to as complex post-traumatic stress disorder (cPTSD; based on the International Classification of Diseases 11th (ICD-11) 20 ). Both cPTSD and the more widely recognised PTSD represent severe psychopathological conditions that substantially compromise individual functioning and psychosocial well-being. Symptom clusters commonly include intrusive recollections such as flashbacks and distressing thoughts, avoidance of trauma-related stimuli, and hyperarousal. In cases of cPTSD, these are often compounded by affective dysregulation (e.g., emotional instability, irritability, or anger outbursts), disturbances in self-perception, and dissociative experiences. This more complex form of trauma response typically results from prolonged exposure to traumatic environments, wherein the traumatic experience becomes integrated into the individual's identity and fundamentally alters self-concept and interpersonal functioning 21 . The next part of our research focuses on the descendants of the survivors – the second generation (G2). Interest in researching the effects of war on the next generation was sparked by vague but consistent symptoms in the descendants of Holocaust survivors 22 – 25 . With further studies, transgenerational transmission became a proven phenomenon, when it was shown that the stress experienced by parents can also affect offspring who were not directly exposed to trauma 26 . The mechanism of transgenerational transmission can occur at several levels (social transmission, epigenetic, prenatal) or their combinations 27 . Unlike animal models, where it is possible to control the conditions of the research 28 , human studies offer only very limited possibilities to determine how and in whom transmission can occur. A study of Kosovar families 11 years after the war found a connection between paternal PTSD, anxiety and depressive symptoms, and depressive symptoms in offspring 29 . In our current work, we must take into account the possible influence of an unfavourable post-war environment. 29 Although the conflict in the former Yugoslavia affected the lives of millions of people, little attention has been paid to it, at least in terms of neuroimaging. The aim of our study is to test whether the impact of the war is detectable after almost three decades, whether and how it affects survivors, and whether it has been transmitted to the next generation growing up in an adverse post-war environment. Results In G1, reduced grey matter volume was observed in the Crus II region of the cerebellum, as well as in the parahippocampal gyrus, fusiform gyrus, middle temporal gyrus, and postcentral gyrus compared to control group (CG). Psychological assessments indicated elevated symptoms of post-traumatic stress (PCL-5), decreased levels of life satisfaction (SWSL), and increased post-traumatic growth indicators (PTGI) in G1 compared to CG. A significant increase in PTGI was found in G2 as compared to CG; no significant differences were found in grey matter volume, post-traumatic stress symptoms, or life satisfaction. MRI Voxel-based morphometry (VBM) revealed a significant GM volume reduction in G1 as compared to control group relevant to G1 (CG1), as shown in Fig. 1 and Table 1. Table 1 Six significant clusters after correction (on each row), where the grey matter volume in G1 is smaller than in CG1 Laterality Structures p-corrected Peak (MNI, mm) Volume L Fusiform ParaHippocampal Cerebellum 4 5 0.0334 -21.5;-30.5;-14.5 1.986 cm3 Temporal Mid 0.0045 -63.5;-21.5;-2.5 3.267 cm3 Occipital Inf 0.0018 -46.5;-79.5;-3.5 1.093 cm3 L and R Cerebellum Crus2 < 0.0001 1.5;-89.5;-31.5 4.22 cm3 R ParaHippocampal Fusiform 0.0015 16.5;-29.5;-7.5 2.802 cm3 Postcentral Precentral 0.0026 54.5;-5.5;29.5 3.054 cm3 Psychological questionnaires The G1 and CG1 groups were compared based on their responses to psychological questionnaires. Significant differences were found in the PTGI, PCL-5, and SWSL questionnaires. G1 had higher scores across all questionnaires. The results are summarised in Table 2. Table 2: Comparison of psychological questionnaire results between survivors (G1 group) and relevant control group Variable G1 Median (IQR) CG1 Median (IQR) p-value PTGI 56 (18) 39 (26) < 0.001 PCL-5 19 (16) 13 (20) 0.009 SWLS 23 (8) 26 (9) 0.040 Brief COPE 67 (12) 66 (10) 0.930 MSPSS 67 (20) 69 (16) 0.4670 The G2 and CG2 groups were compared based on their responses to the psychological questionnaires. Significant differences were found in the PTGI questionnaire. G2 had higher scores than CG2. The results are summarised in Table 3. Table 3 Comparison of psychological questionnaire results between children of survivors (G2 group) and relevant control group Variable G2 Median (IQR) CG2 Median (IQR) p-value PTGI 59 (30) 39 (25) 0.004 PCL-5 19 (17) 20 (25) 0.673 SWLS 23 (10) 26 (7) 0.151 Brief COPE 70 (14) 70 (13) 0.693 MSPSS 72 (15) 70 (12) 0.440 Self-report & Interview questions Survivors of conflicts in the former Yugoslavia (G1 group) In a self-assessment questionnaire, participants rated their satisfaction with their personal and professional lives so far. Overall, they expressed a high level of satisfaction in both areas: 78% of participants were satisfied with their personal lives (Self-report - life) and 80% with their careers (Self-report - career); the results are shown in Graph 1. For the interview question Was the war the hardest thing you have ever experienced in your life? , 53% said no, and 40% said yes. For the question Do you think you coped well with the war? , 77% said yes; however, some elaborated on their experiences with specific triggers that induce a sense of threat. These triggers include the sounds of sirens, ambulances, planes, and similar noises. Additionally, many emphasised that the feeling of threat was heightened by the outbreak of Russian aggression against Ukraine in 2022 and by the uncertainty that accompanied the onset of the Covid-19 pandemic in 2020. The answers are shown in Graph 1 ( War as the hardest thing , Coping with the war ). Children of survivors (G2 group) In the self-report questionnaires (Self-report – life), 71% of G2 were satisfied with their personal life; the overall results are shown in Graph 2. Half of the participants answered yes or rather yes to the interview question Do you think the war had an effect on the way your parents raised you? (Influence on parenting). Several participants expanded this question to include their specific experience of how they had been affected by the war; responses included: sensitivity to the war, pride in parents and country, preoccupation with the topic of the war, fear for loved ones, over-vigilance or caution. To the question Do you think the war had an impact on you, even though you did not experience it directly? (indirect influence of the war), 82% answered yes/rather yes. Several participants expanded on the question by sharing their experiences. Most of them mentioned overprotection by their parents and trying to make sure they were well provided for as children and prepared for emergencies, sometimes even strict upbringing; increased vigilance towards social phenomena; emphasis on not taking what one has for granted; talking about the war – dividing the time before and after the war, but in most cases avoiding overly personal memories. The results for G2 are shown in Graph 2. Discussion In this study, we investigated the long-term effects of war-related stress on structural changes in the brain’s grey matter and in psychological aspects in two generations of people who were directly or indirectly affected by the war in the countries of the former Yugoslavia. Survivors of conflicts in the former Yugoslavia (G1 group) In this study, the most significant finding of the voxel-based morphometry analysis in G1 was a reduction in grey matter volume in the left cerebellar Crus II. Although the cerebellum has historically been associated primarily with motor functions 30 , recent research has confirmed its involvement in non-motor processes, including cognition, emotion, social cognition, and autobiographical memory 31 , 32 . The posterior lobe structure Crus II, part of the cerebellum’s phylogenetically newer regions, has been linked to social mentalisation, expression of self-related emotions, and the processing of autobiographical memory 33 , 34 . A recent meta-analysis identified a consistent association between PTSD and reduced volume in Crus II, with symptom severity showing a stronger correlation than diagnosis alone 35 . Functional MRI studies have also shown increased activation of Crus II during the recall of sad and negative autobiographical memories 33 . The posterior cerebellum is anatomically connected to paralimbic areas and contributes to the integration of emotion and behaviour 35 . A reduction in volume was also found in the right and left parahippocampus, part of the paralimbic system. The parahippocampal gyrus (PHG), located on the ventromedial surface of the temporal lobe and closely associated with the hippocampus, plays a role in emotional processing and memory consolidation 36 . Nardo and his team focused their study on grey matter concentration; they found a reduction in density in the PHG (and in the posterior cingulate and insular cortex) in PTSD subjects. They suggest that the altered function of PHG may lead to memory impairments seen in PTSD or impaired processing of emotional stimuli and integration of traumatic memories 37 . Also, reduction in the volume of this structure has been found in patients with PTSD 38 . In another work 39 , the authors argue that disrupted parahippocampal-prefrontal coupling may result in altered memory suppression in patients with PTSD 39 . A reduction in grey matter was also observed in the middle temporal gyrus (MTG), which is involved in cognition and memory processing, particularly in the retrieval and suppression of emotional memories, including PTSD flashbacks 40 . Volume reduction in the left MTG has been associated with impaired retrieval of autobiographical memories in PTSD 41 . The fusiform gyrus (FG), known for its role in face and body recognition and complex visual processing, is also involved in memory, emotional perception, and threat detection. Reduced volume in this region has been linked to deficits in visual and emotional processing of social stimuli in PTSD, potentially contributing to symptoms such as hypervigilance and social dysfunction 38 , 42 , 43 . Although the postcentral gyrus, which is primarily responsible for somatosensory processing, is not usually examined in research on psychological trauma, a study of war veterans with PTSD was conducted to examine somatosensory responses to non-threatening stimuli. Veterans with PTSD did not show typical responses to touch in the postcentral gyrus, superior parietal region, and right prefrontal cortex.Unlike participants with PTSD responded differently to non-threatening tactile stimuli This result suggests that PTSD manifests itself not only through psychological symptoms but also through somatosensory symptoms, such as altered responses to tactile stimuli 44 . The finding that all regions in which we found volume reductions in G1 were associated with post-traumatic stress disorder or its symptoms corresponds with the results of the PCL-5 questionnaire, which revealed higher symptoms of persistent post-traumatic stress in G1. The threshold of 31 points or more 45 , on the basis of which a preliminary diagnosis of PTSD can be made, was exceeded in approximately half of the subjects in group G1. Therefore, while it cannot be conclusively stated that individuals in G1 meet the full diagnostic criteria for PTSD, they do exhibit pronounced symptoms indicative of post-traumatic stress. Furthermore, we found lower satisfaction with life (SWSL) in G1 as compared to CG. Similar results were found in an earlier study that examined individuals affected by the conflict in the former Yugoslavia 10 years later. These individuals still showed symptoms of post-traumatic stress decreased quality of life 10 years after the conflict 46 . Our results extend these findings to a broader time horizon. From our previous work examining Holocaust survivors 7 , we know that increased stress symptoms and lower well-being can persist over the long term and even throughout an entire lifespan. However, the same study found that post-traumatic growth also persists, as is the case with G1 in the current study. The coexistence of post-traumatic growth and stress is not an uncommon phenomenon. Some studies have explored the possibility that PTG is not a completed event based on an experienced event, but a process of seeking to cope with stress 47 . Given that the reduction in grey matter volume in G1 occurred in the Crus II area, which is demonstrably involved in the processing of negative memories and social mentalisation, one plausible explanation for this reduction is the suppression of negative experiences, which may function as a coping mechanism in G1. This hypothesis is indirectly supported by interviews from our G2 participants, who are the children of war survivors, even though they are not direct descendants of our G1 participants. In interviews, G2 often reported that discussions of the war were common in their households. However, these conversations typically lacked deep introspection regarding the emotional experiences of their parents. Empirical studies suggest that while thought suppression or avoidance may offer short-term relief, it is associated with the persistence of PTSD symptoms in the long term 48 . This finding is consistent with the psychological questionnaires used in this study, which indicate that G1 participants exhibit higher levels of PTSD symptoms and lower life satisfaction. However, results from the PTGI suggest that, despite these challenges, G1 participants also experience post-traumatic growth, reporting positive evaluations of their personal and professional lives. Many have achieved above-average incomes, high levels of education, and have successfully integrated into countries other than their homeland. Although suppression is generally considered a maladaptive long-term coping strategy, it does not preclude the possibility of resilience. In their research on resilience in post-war Southeastern Europe, Kelmendi and Hamby identified emotional self-regulation as a prominent trauma coping mechanism characteristic of this region 49 . In G1, the coping mechanism appears to involve conscious or unconscious suppression of negative autobiographical memories, leading to the creation of a “protective shell” that allows them to cope with trauma and live a full life, without completely eliminating their emotional scars. Children of survivors (G2 group) The second generation in this study consisted of people who were born after the end of the war to parents affected by the war (in Bosnia and Herzegovina or in Croatia) or the bombing in Serbia in 1999. Most participants grew up in the post-war countries of the former Yugoslavia, some were born elsewhere, and a few participants were born and spent their childhood in the Czech Republic, although they continued to have a relationship with their parents’ countries of origin. In the interviews, the participants reported that the topic of war is still alive. It was repeatedly mentioned that for some families, the war is a constant part of almost every conversation, and time is split into pre-war and post-war periods, with parents often talking about what happened in terms of their perspective of everyday life but sharing little about their inner experiences and feelings. We found no differences between the brain structures of the G2 and their peers in CG2. Neither were differences found in the PCL-5 and SWSL questionnaires: participants do not show differences in persistent symptoms of post-traumatic stress nor feel lower satisfaction with life compared to the control group. Like G1, G2 do show higher post-traumatic growth than CG. However, participants from G2 did not have a significant difference in PCL-5, nor did they exceed the cut-off threshold for PCL-5 in greater numbers (only three subjects had scores above 31). Thus, it cannot be said that this group carried persistent post-traumatic stress. The absence of post-traumatic stress and the presence of post-traumatic growth are not mutually exclusive. Although in most interviews, with the exception of two participants, participants in G2 rated the war as having had an impact on them, a possible explanation is that they have successfully integrated the difficulties they experienced into their lives, or that PTG has served as an adaptation process, as we indicated above for G1 47 . Based on our recent research, we found that the second generation differed in some ways from the non-war-affected control group, but we could not determine whether this was due to the post-war environment 50 , family and collective history 51 , transgenerational transmission, or a combination of these factors. It seems that these questions will be the subject of new research in the future 52 . Limitations Our group cannot represent an independent cross-section of the Balkan population. In our research, we only worked with individuals who volunteered for the study; their motivation to participate itself may have biased the sample. Furthermore, these people have an above-average socioeconomic background; G1 participants had lived in a country other than their home country for a long time. G1 participants had experienced war in many forms and in different countries of the former Yugoslavia (Bosnia and Herzegovina, Croatia, Serbia). It is not a perfectly homogeneous group, but on the other hand, it is a model sample of people affected by war in various forms. Our two stress groups, G1 and G2, are not parents and children. Therefore, we cannot observe direct transmission from parents to offspring. Conclusion Individuals affected by the war in the former Yugoslavia exhibit reduced grey matter volume in brain structures associated with post-traumatic stress disorder (PTSD) symptoms and autobiographical memory processing. These individuals continue to experience persistent symptoms of post-traumatic stress and report lower life satisfaction compared to control participants. At the same time, they demonstrate higher levels of post-traumatic growth and, in self-reports, assess their lives positively. This pattern suggests the presence of a specific, albeit imperfect, coping mechanism that enables a more reactive response to perceived or actual threats, while still allowing for the pursuit of a fulfilling life. In contrast, the second generation, raised in a challenging post-war environment shaped by a pervasive war narrative, do not exhibit structural brain changes or significant differences in life satisfaction and stress symptoms compared to their peers in the control group. Although they do not show persistent symptoms of post-traumatic stress, they tend to perceive that the difficulties caused by the post-war environment have a negative impact on their lives. At the same time, they report increased post-traumatic growth, suggesting that they may have effectively integrated these challenges into their lives or that post-traumatic growth serves as an adaptive ongoing mechanism supporting their ability to cope with adversity. Finally, these findings highlight the need for stress research to consider the role of the cerebellum, particularly Crus II, which has long been overlooked in studies of non-motor brain functions Methods and analysis Research The research was conducted between 2022 and 2025 at the Central European Institute of Technology—Research Centre of Masaryk University in Brno. It was approved by the Ethics Committee of Masaryk University in accordance with the Declaration of Helsinki. The ethics committee’s approval code number is EKV-2021-076. Informed consent was obtained from each participant. This project represents an interdisciplinary research effort examining the psychological, physiological, and neuroimaging correlates of war-related stress in civilian populations. The current study specifically focuses on data pertinent to the assessment of brain structure, utilising structural MRI findings alongside psychological questionnaires and interview data. Recruitment Volunteers were recruited between 2022 and 2024 through media reports, university information channels, social networks, and a lecture at the Lastavica club, which brings together the Balkan diaspora in the Czech Republic. Participants We investigated participants who, as civilians, survived the war in former Yugoslavia and second-generation survivors – children of war survivors. The group of participants we categorised as the first generation (G1) was composed of people (n = 45) who experienced multiple war events and were under life-threatening stress during war activity in then-Yugoslavia in 1991–1995 (Bosna and Herzegovina, Croatia) or experienced NATO bombing in Serbia in 1999, or, in some cases, a combination of these events. To the question, How did you survive the war? What did you experience during the war? participants provided a variety of responses, including: Shelling and bombing - Hiding in a cellar, shelter, or other hiding places Forced abandonment or destruction of homes War-related injuries - Death or injury of loved ones Witnessing the death or injury of others Separation from family and loved ones Fear for the safety of loved ones Lack of food, medicine, water, and electricity Poverty Discriminatory and humiliating behaviour by soldiers Involuntary service in the military (none of the participants were professional soldiers) Inconveniences associated with forced emigration, including bullying or rejection, loss of social capital, and financial difficulties. After the experience of the conflict, these people emigrated from their home country directly to the Czech Republic or to other countries and then to the Czech Republic, where they currently live. The second generation (G2) contains people who are descendants of war survivors (n = 28) and were not exposed to war-threatening stress but grew up in a post-war environment. In addition to five persons of Czech nationality who were born in the Czech Republic to parents who had emigrated to the Czech Republic, there were participants of Bosnian, Croatian, Serbian and Montenegrin nationality who were temporarily in the Czech Republic as part of a study programme. G1 and G2 are not family members (with a few exceptions). G2 participants are not direct descendants of G1. The control group (CG) consisted of Czechs or Slovaks living in Czechia who had no war experienc; the group were divided into two subgroups corresponding to G1 and G2, we called the subgroups CG1 and CG2 (n = 73; 45 + 28). Exclusion criteria Brain impairment (brain injuries or diseases, tumours, neurodegenerative diseases), severe psychiatric disorders*(e.g. psychosis), significant cognitive decline. Furthermore, participants with technical artefacts or excessive movement artefacts in the MR images were excluded. * Due to the nature of the research, individuals who were previously diagnosed with post-traumatic stress disorder or believed they might be diagnosed with it were retained in the sample. The stress groups and their assigned control groups did not differ significantly in age, sex, or education. Income was significantly higher in G1 than in CG1. The education and income of G2 and CG2 were not assessed because most participants were students. Age was compared using the two-sample t-test; sex, education, and income were compared using the Fisher exact test (Table 4). Table 4 Demographic data of the research population G1 CG1 p-value Count 45 45 - Age 38 ± 9 37 ± 10 0.5870 Sex 23 M / 22 F 20 M / 25 F 0.6732 University education 29 Y / 16 N 32 Y / 13 F 0.6523 Income 30 H / 15 L 11 H / 34 L 0.0001 G2 CG2 p-value Count 28 28 - Age 25 ± 3 24 ± 3 0.3255 Sex 9 M / 19 F 11 M / 18 F 0.7828 MRI MR examinations were performed on a 3T scanner Siemens Prisma using a 64-channel head coil. The MRI protocol for voxel-based morphometry included 3D T1-weighted magnetisation prepared rapid gradient echo (MPRAGE) sequence with TR = 2.3 s, TE = 2.33 ms, TI = 0.9 s, FA = 8°, isometric voxel size 1 mm in FOV 224 × 224 mm and 240 slices. Anatomical MRI data were analysed using SPM12 (www.fil.ion.ucl.ac.uk) and CAT12 toolbox (www.neuro.uni-jena.de/cat) running in Matlab R2020a. The standard recommended pipeline was used for segmentation, in short: high resolution data were segmented into grey matter (GM) using the SPM Tissue Probability Map (TPM) and registered into common MNI space using shooting template IXI555_MNI152_GS. Finally, spatially normalised and modulated GM maps were smoothed with 6 mm FWHM isotropic Gaussian kernel. Group-level statistics for stress effects were computed using a second-level model in SPM12. Modulated GM images were corrected for total intracranial volume and subsequently analysed. A two-sample t-test compared GMV maps between the stress groups (G1, G2) and control groups (CG1, CG2); sex and age were included as nuisance variables. Resultant t-statistic maps were initially thresholded at a P value of < 0.001 uncorrected and then only significant clusters at P < 0.05 FWE cluster level were picked. Psychological questionnaires & Self-report & Interview Participants completed a set of psychological questionnaires (Table 5), a self-assessment, and a semi-structured interview, allowing them to answer questions briefly or provide more detailed responses if they wished. Table 5 Psychological questionnaires PTGI Post-Traumatic Growth Inventory Assesses positive outcomes of experiencing stressful life events. The Inventory contains 21 items and measures 5 factors: New Possibilities, Relating to Others, Personal Strength, Spiritual Change, and Appreciation of Life. Participants were asked to mark the degree of perceived change (0 = I did not experience this change as a result of my crisis to 5 = I experienced this change to a very great degree). The score range is from 0 to 105, with higher scores indicative of greater post-traumatic growth 53 . PCL-5 The PTSD Check List for DSM-5 A 20-item self-report instrument for measuring symptoms of PTSD based on DSM-5. Respondents are asked to consider a “list of problems and complaints that people sometimes have in response to stressful experiences” that is supposed to rate how much they “have been bothered by each problem in the past month”. Every item is scaled from 0 (“not at all”) to 4 (“extremely”). Completing the inventory takes approximately 5–10 minutes. The PLC-5 questionnaire can be evaluated in several ways. A cut-off score between 31 and 33 items is used for a provisional diagnosis of PTSD 45 . SWSL Satisfaction with Life Scale A 5-item self-report instrument. Respondent answers are marked on Likert scale ranging from 1 (“strongly disagree”) to 7 (“strongly agree”), and are presented in raw scores, with a total score range from 5 to 35. The SWLS focuses on cognitive aspects of life satisfaction rather than emotional; moreover, SWLS does not focus on specific domains. SWLS makes it possible to weigh different aspects of life based on personal criteria 54 . MSPSS Multidimensional Scale of Perceived Social Support A 12-item self-report scale. The instrument measures subjectively assessed social support based on three subscales addressing a different source of support: a) family, b) friends and c) significant others. Respondent answers are marked on Likert scale ranging from 1 (“very strongly disagree”) to 7 (“very strongly agree”) 55 Brief COPE A 28-item self-report inventory focusing on effective and ineffective ways to cope with stressful events. It is useful tool for detecting emotional responses to serious events e.g. health conditions, financial stress, mental illness, injuries and more. Respondent answers were marked on scale ranging from 1 (“I haven't been doing this at all”) to 4 (“I've been doing this a lot”) 56 . The results of the psychological questionnaires were summarised using the median, as well as the Interquartile range (IQR). To test for statistical differences, the non-parametric Mann-Whitney U test was employed. The significance level for all statistical tests was set at p < 0.05. Self-report & interview All participants completed a short self-assessment of their life and work achievements to date. Satisfaction with life and career was rated on a scale of yes, rather yes, somewhere in between, rather no, no. Participants from G2 did not rate career success as they were predominantly students. Stress groups (G1 and G2) additionally provided information on war experiences affecting themselves or their families through semi-structured interviews, forming the basis for a separate psychological study. For the current text, we report only demographic data, life satisfaction ratings, and war-related experiences. G1 participants were asked whether war was the worst event of their life, and how they coped; G2 participants were asked whether they felt that the war had affected them indirectly and whether the war had affected their parents’ parenting style. Abbreviations G1 - First Generation G2 - Second Generation CG - Control Group CG1 - Control Group for G1 CG2 - Control Group for G2 MRI - Magnetic Resonance Imaging PTSD - Post-Traumatic Stress Disorder cPTSD - Complex Post-Traumatic Stress Disorder ICD-11 - International Classification of Diseases, 11th Revision PTGI - Post-Traumatic Growth Inventory PCL-5 - PTSD Checklist for DSM-5 SWSL - Satisfaction with Life Scale MSPSS - Multidimensional Scale of Perceived Social Support Brief-COPE - Brief Coping Orientation to Problems Experienced Inventory Declarations Funding This study was supported by a grant from the Ministry of Health of the Czech Republic, grant no. NU22-04-00661. Data availability The datasets used and/or analysed during the current study available from the corresponding author on reasonable request. The authors assert that all procedures contributing to this work comply with the ethical standards of the relevant national and institutional committees on human experimentation and with the Helsinki Declaration of 1975, as revised in 2008. Acknowledgement We thank Anne Johnson for grammatical assistance. We acknowledge the core facility MAFIL of CEITEC MU. We acknowledge the core facility MAFIL supported by the Czech-BioImaging large RI project (LM2023050 funded by MEYS CR), part of the Euro-BioImaging (www.eurobioimaging.eu) ALM and Multimodal Imaging Node (Brno, CZ), for their support with obtaining scientific data presented in this paper. Author information Authors and Affiliations Central European Institute of Technology (CEITEC), Brain and Mind Research Programme, Masaryk University, Brno, Czechia Monika Fňašková, Pavel Říha, Marek Preiss, David Ulčák, Markéta Nečasová, Nikola Wolframová, Martin Lamoš & Ivan Rektor National Institute of Mental Health (Czechia), Prague, Czechia Marek Preiss 3. First Department of Neurology, St Anne’s University Hospital and Faculty of Medicine, Masaryk University, Brno, Czechia Monika Fňašková, Pavel Říha, David Ulčák, Martin Lamoš & Ivan Rektor 4. University of New York in Prague Marek Preis 4. Utrecht University Vojtěch Svoboda Contributions M.F. (corresponding author): Project Administration, Investigation, Resources, Data Curation, Writing Original Draft, Visualisation, P.Ř. - Methodology, Software, Formal Analysis, Investigation, Visualisation, M.P. - Conceptualisation, Methodology, Validation, D. U. Investigation, Visualisation, M.N. - Investigation, N.W. Investigation, V. S. Investigation, M.L. Supervision, I.R. - Conceptualisation, Supervision. Corresponding author Correspondence to Monika Fňašková ( [email protected] ) Ethics declarations Competing interests The authors declare no competing interests. Ethics approval statement Ethical approval for this study was obtained from the ethics committee of Masaryk University (approval code EKV-2021-076) on June 24, 2021 References Comtesse, H., Powell, S., Soldo, A., Hagl, M. & Rosner, R. Long-term psychological distress of Bosnian war survivors: an 11-year follow-up of former displaced persons, returnees, and stayers. BMC Psychiatry 19 , 1 (2019). Priebe, S. et al. Psychological Symptoms as Long-Term Consequences of War Experiences. Psychopathology 46 , 45–54 (2013). Ćopić, S. Wife Abuse in the Countries of the Former Yugoslavia. Feminist Review 76 , 46–64 (2004). Jovanović, R. & Bermúdez, Á. The next generation: Nationalism and violence in the narratives of Serbian students on the break-up of Yugoslavia. 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Cerebellum 23 , 2332–2340 (2024). Van Overwalle, F., Ma, Q. & Heleven, E. The posterior crus II cerebellum is specialized for social mentalizing and emotional self-experiences: a meta-analysis. Soc Cogn Affect Neurosci 15 , 905–928 (2020). Huggins, A. A. et al. Smaller total and subregional cerebellar volumes in posttraumatic stress disorder: a mega-analysis by the ENIGMA-PGC PTSD workgroup. Mol Psychiatry 29 , 611–623 (2024). Lew, C. H. & Semendeferi, K. 4.16 - Evolutionary Specializations of the Human Limbic System. in Evolution of Nervous Systems (Second Edition) (ed. Kaas, J. H.) 277–291 (Academic Press, Oxford, 2017). doi:10.1016/B978-0-12-804042-3.00115-9. Nardo, D. et al. Gray matter density in limbic and paralimbic cortices is associated with trauma load and EMDR outcome in PTSD patients. Journal of Psychiatric Research 44 , 477–485 (2010). Kunimatsu, A., Yasaka, K., Akai, H., Kunimatsu, N. & Abe, O. MRI findings in posttraumatic stress disorder. Journal of Magnetic Resonance Imaging 52 , 380–396 (2020). Steward, T., Das, P., Malhi, G. S., Bryant, R. A. & Felmingham, K. L. Dysfunctional coupling of the parahippocampal cortex and inferior frontal gyrus during memory suppression in posttraumatic stress disorder. European Neuropsychopharmacology 41 , 146–151 (2020). Brewin, C. R., Gregory, J. D., Lipton, M. & Burgess, N. Intrusive images in psychological disorders: Characteristics, neural mechanisms, and treatment implications. Psychological Review 117 , 210–232 (2010). Li, L. et al. Grey matter reduction associated with posttraumatic stress disorder and traumatic stress. Neuroscience & Biobehavioral Reviews 43 , 163–172 (2014). Mueller, E. M. & Pizzagalli, D. A. One-year-old fear memories rapidly activate human fusiform gyrus. Social Cognitive and Affective Neuroscience 11 , 308–316 (2016). Weiner, K. S. & Zilles, K. The anatomical and functional specialization of the fusiform gyrus. Neuropsychologia 83 , 48–62 (2016). Badura-Brack, A. S. et al. Decreased somatosensory activity to non-threatening touch in combat veterans with posttraumatic stress disorder. Psychiatry Research: Neuroimaging 233 , 194–200 (2015). Blevins, C. A., Weathers, F. W., Davis, M. T., Witte, T. K. & Domino, J. L. The Posttraumatic Stress Disorder Checklist for DSM-5 (PCL-5): Development and Initial Psychometric Evaluation. J Trauma Stress 28 , 489–498 (2015). Priebe, S. et al. Consequences of Untreated Posttraumatic Stress Disorder Following War in Former Yugoslavia: Morbidity, Subjective Quality of Life, and Care Costs. Croatian Medical Journal 50 , 465–475 (2009). Dekel, S., Ein-Dor, T. & Solomon, Z. Posttraumatic growth and posttraumatic distress: A longitudinal study. Psychological Trauma: Theory, Research, Practice, and Policy 4 , 94–101 (2012). Holmes, E. A., Moulds, M. L. & Kavanagh, D. Memory Suppression in PTSD Treatment? Science 318 , 1722–1722 (2007). Kelmendi, K. & Hamby, S. Resilience After Trauma in Kosovo and Southeastern Europe: A Scoping Review. Trauma, Violence, & Abuse 24 , 2333–2345 (2023). Wasiak, K. Between Trauma and Nostalgia: Second Generation Memory of the War in Bosnia and Herzegovina. Historia i Polityka 53 , 71–83 (2023). Audergon, A. Collective trauma: the nightmare of history. Psychotherapy and Politics International 2 , 16–31 (2004). Sangalang, C. C. The scars of war last for centuries: how we understand collective trauma needs to change. Nature 641 , 589–591 (2025). Tedeschi, R. G. & Calhoun, L. G. The Posttraumatic Growth Inventory: Measuring the positive legacy of trauma. J Trauma Stress 9 , 455–471 (1996). Diener, E., Emmons ,Robert A., Larsen ,Randy J. & and Griffin, S. The Satisfaction With Life Scale. Journal of Personality Assessment 49 , 71–75 (1985). Zimet, G. D., Dahlem ,Nancy W., Zimet ,Sara G. & and Farley, G. K. The Multidimensional Scale of Perceived Social Support. Journal of Personality Assessment 52 , 30–41 (1988). Carver, C. S. You want to measure coping but your protocol’ too long: Consider the brief cope. Int. J. Behav. Med. 4 , 92–100 (1997). Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 17 Mar, 2026 Read the published version in Scientific Reports → Version 1 posted Editorial decision: Revision requested 15 Sep, 2025 Reviews received at journal 25 Aug, 2025 Reviews received at journal 15 Aug, 2025 Reviewers agreed at journal 14 Aug, 2025 Reviewers agreed at journal 06 Aug, 2025 Reviewers invited by journal 30 Jul, 2025 Editor assigned by journal 22 Jul, 2025 Editor invited by journal 07 Jul, 2025 Submission checks completed at journal 07 Jul, 2025 First submitted to journal 02 Jul, 2025 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. 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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-7028508","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":494965492,"identity":"36edf5c4-a12c-4052-b840-c5b13fd34996","order_by":0,"name":"Monika 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University","correspondingAuthor":false,"prefix":"","firstName":"Martin","middleName":"","lastName":"Lamoš","suffix":""},{"id":494965500,"identity":"1a60f611-2306-47e3-a635-842a39013884","order_by":8,"name":"Ivan Rektor","email":"","orcid":"","institution":"Central European Institute of Technology (CEITEC), Masaryk University","correspondingAuthor":false,"prefix":"","firstName":"Ivan","middleName":"","lastName":"Rektor","suffix":""}],"badges":[],"createdAt":"2025-07-02 10:53:24","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7028508/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7028508/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41598-026-44241-w","type":"published","date":"2026-03-17T15:59:43+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":88310915,"identity":"d6773a8e-7e50-4559-80a1-56a111df8c18","added_by":"auto","created_at":"2025-08-05 06:54:15","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":277944,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eComparison of grey matter volume between survivors (G1 group) and relevant control group\u003c/em\u003e\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-7028508/v1/155c78211d74f337c1ef6482.png"},{"id":105223442,"identity":"2c9daac8-83ec-414f-ae21-9e0d565c78b0","added_by":"auto","created_at":"2026-03-23 16:06:46","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1324414,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7028508/v1/f2cf0f89-51d3-4469-aec8-a41f7c1060ff.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Long-term impact of war stress in civilians from the former Yugoslavia: A two-generation neurostructural and psychological study","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe conflict in the former Yugoslavia, which lasted from the 1990s to the beginning of the new millennium, was one of the worst and most extensive in Europe since World War II. The wars that successively affected Slovenia, Croatia, Bosnia and Herzegovina, and Kosovo beginning in 1991, and the NATO bombing of Serbia and Montenegro in 1999 in response to the conflict in Kosovo, left the population affected by many traumatic experiences including war injuries, death of loved ones, separation of families, involuntary displacement from their homes, sexual violence, and torture \u003csup\u003e\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e,\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e\u003c/sup\u003e. Post-war development in these countries devastated by war or bombing has been difficult. Psychological help has not been routinely available, and untreated trauma has often manifested itself in alcohol abuse or violence \u003csup\u003e\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e\u003c/sup\u003e. The impact of poverty and harsh conditions, and the daily narrative of the war, has taken a toll on the next generation \u003csup\u003e\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e,\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eThis study investigates the long-term effects of war-related stress on civilian populations (G1) and their descendants (G2) from the countries of the former Yugoslavia. Utilising structural magnetic resonance imaging, a psychological survey, and interviews, we explore whether and how the impacts of war have influenced their current lives.\u003c/p\u003e\u003cp\u003eNeuroimaging studies investigating the effects of traumatic psychological stress have consistently demonstrated alterations in brain structure and function, including reduced grey matter volume, often associated with the development of PTSD \u003csup\u003e\u003cspan additionalcitationids=\"CR7 CR8\" citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e. Although the hippocampus, amygdala, and prefrontal cortex are commonly identified as the primary stress-sensitive structures, the results vary and many other regions undergo structural or functional changes due to stress \u003csup\u003e\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e,\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e\u003c/sup\u003e. In addition to the timing, intensity, and duration of psychological stress, one of the main factors affecting the brain appears to be the nature of the traumatic experience \u003csup\u003e\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e\u003c/sup\u003e. Studies dealing with the effect of war trauma on the alteration of brain structures in non-military personnel are scarce; most studies have been conducted on soldiers or war veterans \u003csup\u003e\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e,\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u003c/sup\u003e. Even though research on civilians is more demanding due to more diverse war experiences, non-existent or inadequate prior medical records, financial demands of research or non-existent research facilities in a war-torn country, and many other reasons, it is an important topic that should be given adequate attention, because unexplored and untreated trauma may have far-reaching effects on individuals and entire societies. In the context of the war in the former Yugoslavia, we found only one MRI paper focused on brain structure volumes conducted on a group of Croatian war veterans who were not professional soldiers. The authors found a significant reduction in right hippocampal volume and a non-significant reduction in left hippocampal volume in this group \u003csup\u003e\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eThe psychological consequences of war include depressive and other mood disorders, anxiety, paranoid ideations, and post-traumatic stress disorder (PTSD) \u003csup\u003e\u003cspan additionalcitationids=\"CR17\" citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e\u003c/sup\u003e. PTSD is generally considered to be an illness caused by a traumatic experience. It has been reported that in the general population, PTSD arises as a result of trauma in about 5\u0026ndash;10% of cases; in the case of war stress, the incidence of PTSD roughly doubles, and as many as 26% of persons who have remained in a war-afflicted region have been found to have the condition \u003csup\u003e\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e\u003c/sup\u003e. The condition and its symptomatology are frequently examined in relation to experiences of armed conflict, with documented impacts on both military personnel and civilian populations. In contexts of sustained or repeated exposure to stress characteristic of war and displacement, a more pervasive psychological response may develop, clinically referred to as complex post-traumatic stress disorder (cPTSD; based on the International Classification of Diseases 11th (ICD-11) \u003csup\u003e\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e\u003c/sup\u003e).\u003c/p\u003e\u003cp\u003eBoth cPTSD and the more widely recognised PTSD represent severe psychopathological conditions that substantially compromise individual functioning and psychosocial well-being. Symptom clusters commonly include intrusive recollections such as flashbacks and distressing thoughts, avoidance of trauma-related stimuli, and hyperarousal. In cases of cPTSD, these are often compounded by affective dysregulation (e.g., emotional instability, irritability, or anger outbursts), disturbances in self-perception, and dissociative experiences. This more complex form of trauma response typically results from prolonged exposure to traumatic environments, wherein the traumatic experience becomes integrated into the individual's identity and fundamentally alters self-concept and interpersonal functioning \u003csup\u003e\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eThe next part of our research focuses on the descendants of the survivors \u0026ndash; the second generation (G2). Interest in researching the effects of war on the next generation was sparked by vague but consistent symptoms in the descendants of Holocaust survivors \u003csup\u003e\u003cspan additionalcitationids=\"CR23 CR24\" citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e\u003c/sup\u003e. With further studies, transgenerational transmission became a proven phenomenon, when it was shown that the stress experienced by parents can also affect offspring who were not directly exposed to trauma \u003csup\u003e\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e\u003c/sup\u003e. The mechanism of transgenerational transmission can occur at several levels (social transmission, epigenetic, prenatal) or their combinations \u003csup\u003e\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e\u003c/sup\u003e. Unlike animal models, where it is possible to control the conditions of the research \u003csup\u003e\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e\u003c/sup\u003e, human studies offer only very limited possibilities to determine how and in whom transmission can occur. A study of Kosovar families 11 years after the war found a connection between paternal PTSD, anxiety and depressive symptoms, and depressive symptoms in offspring \u003csup\u003e\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u003c/sup\u003e. In our current work, we must take into account the possible influence of an unfavourable post-war environment. \u003csup\u003e\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u003c/sup\u003e\u003c/p\u003e\u003cp\u003eAlthough the conflict in the former Yugoslavia affected the lives of millions of people, little attention has been paid to it, at least in terms of neuroimaging. The aim of our study is to test whether the impact of the war is detectable after almost three decades, whether and how it affects survivors, and whether it has been transmitted to the next generation growing up in an adverse post-war environment.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eIn G1, reduced grey matter volume was observed in the Crus II region of the cerebellum, as well as in the parahippocampal gyrus, fusiform gyrus, middle temporal gyrus, and postcentral gyrus compared to control group (CG). Psychological assessments indicated elevated symptoms of post-traumatic stress (PCL-5), decreased levels of life satisfaction (SWSL), and increased post-traumatic growth indicators (PTGI) in G1 compared to CG.\u003c/p\u003e\n\u003cp\u003eA significant increase in PTGI was found in G2 as compared to CG; no significant differences were found in grey matter volume, post-traumatic stress symptoms, or life satisfaction.\u003c/p\u003e\n\u003cp\u003eMRI\u003c/p\u003e\n\u003cp\u003eVoxel-based morphometry (VBM) revealed a significant GM volume reduction in G1 as compared to control group relevant to G1 (CG1), as shown in Fig. 1 and Table 1.\u003c/p\u003e\n\u003cdiv\u003e\n \u003ctable id=\"Tab1\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv\u003eTable 1\u003c/div\u003e\n \u003cdiv\u003e\n \u003cp\u003eSix significant clusters after correction (on each row), where the grey matter volume in G1 is smaller than in CG1\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eLaterality\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eStructures\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ep-corrected\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePeak (MNI, mm)\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eVolume\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"3\"\u003e\n \u003cp\u003eL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eFusiform\u003c/p\u003e\n \u003cp\u003eParaHippocampal\u003c/p\u003e\n \u003cp\u003eCerebellum 4 5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.0334\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e-21.5;-30.5;-14.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1.986 cm3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eTemporal Mid\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.0045\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e-63.5;-21.5;-2.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3.267 cm3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eOccipital Inf\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.0018\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e-46.5;-79.5;-3.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1.093 cm3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eL and R\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eCerebellum Crus2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026lt; 0.0001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.5;-89.5;-31.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.22 cm3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eR\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eParaHippocampal\u003c/p\u003e\n \u003cp\u003eFusiform\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.0015\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e16.5;-29.5;-7.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.802 cm3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePostcentral\u003c/p\u003e\n \u003cp\u003ePrecentral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.0026\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e54.5;-5.5;29.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3.054 cm3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003ePsychological questionnaires\u003c/p\u003e\n\u003cp\u003eThe G1 and CG1 groups were compared based on their responses to psychological questionnaires. Significant differences were found in the PTGI, PCL-5, and SWSL questionnaires. G1 had higher scores across all questionnaires. The results are summarised in Table 2.\u003c/p\u003e\n\u003cdiv\u003e\n \u003ctable id=\"Tab2\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv\u003e\u003cem\u003eTable 2: Comparison of psychological questionnaire results between survivors (G1 group) and relevant control group\u003c/em\u003e\u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eVariable\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eG1 Median (IQR)\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCG1 Median (IQR)\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ep-value\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003ePTGI\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e56 (18)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e39 (26)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026lt; 0.001\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003ePCL-5\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e19 (16)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e13 (20)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u003cstrong\u003e0.009\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eSWLS\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23 (8)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e26 (9)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u003cstrong\u003e0.040\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eBrief COPE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e67 (12)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e66 (10)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.930\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eMSPSS\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e67 (20)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e69 (16)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.4670\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003eThe G2 and CG2 groups were compared based on their responses to the psychological questionnaires. Significant differences were found in the PTGI questionnaire. G2 had higher scores than CG2. The results are summarised in Table 3.\u003c/p\u003e\n\u003cdiv\u003e\n \u003ctable id=\"Tab3\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv\u003eTable 3\u003c/div\u003e\n \u003cdiv\u003e\n \u003cp\u003eComparison of psychological questionnaire results between children of survivors (G2 group) and relevant control group\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eVariable\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eG2 Median (IQR)\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCG2 Median (IQR)\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ep-value\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003ePTGI\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e59 (30)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e39 (25)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u003cstrong\u003e0.004\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003ePCL-5\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e19 (17)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20 (25)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.673\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eSWLS\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23 (10)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e26 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.151\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eBrief COPE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e70 (14)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e70 (13)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.693\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eMSPSS\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e72 (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e70 (12)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.440\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003eSelf-report \u0026amp; Interview questions\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eSurvivors of conflicts in the former Yugoslavia (G1 group)\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eIn a self-assessment questionnaire, participants rated their satisfaction with their personal and professional lives so far. Overall, they expressed a high level of satisfaction in both areas: 78% of participants were satisfied with their personal lives (Self-report - life) and 80% with their careers (Self-report - career); the results are shown in Graph 1. For the interview question \u003cem\u003eWas the war the hardest thing you have ever experienced in your life?\u003c/em\u003e, 53% said no, and 40% said yes. For the question \u003cem\u003eDo you think you coped well with the war?\u003c/em\u003e, 77% said yes; however, some elaborated on their experiences with specific triggers that induce a sense of threat. These triggers include the sounds of sirens, ambulances, planes, and similar noises. Additionally, many emphasised that the feeling of threat was heightened by the outbreak of Russian aggression against Ukraine in 2022 and by the uncertainty that accompanied the onset of the Covid-19 pandemic in 2020. The answers are shown in Graph 1 (\u003cem\u003eWar as the hardest thing\u003c/em\u003e, \u003cem\u003eCoping with the war\u003c/em\u003e).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eChildren of survivors (G2 group)\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eIn the self-report questionnaires (Self-report – life), 71% of G2 were satisfied with their personal life; the overall results are shown in Graph 2.\u003c/p\u003e\n\u003cp\u003eHalf of the participants answered yes or rather yes to the interview question \u003cem\u003eDo you think the war had an effect on the way your parents raised you?\u003c/em\u003e (Influence on parenting). Several participants expanded this question to include their specific experience of how they had been affected by the war; responses included: sensitivity to the war, pride in parents and country, preoccupation with the topic of the war, fear for loved ones, over-vigilance or caution. To the question \u003cem\u003eDo you think the war had an impact on you, even though you did not experience it directly?\u003c/em\u003e (indirect influence of the war), 82% answered yes/rather yes. Several participants expanded on the question by sharing their experiences. Most of them mentioned overprotection by their parents and trying to make sure they were well provided for as children and prepared for emergencies, sometimes even strict upbringing; increased vigilance towards social phenomena; emphasis on not taking what one has for granted; talking about the war – dividing the time before and after the war, but in most cases avoiding overly personal memories. The results for G2 are shown in Graph 2.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn this study, we investigated the long-term effects of war-related stress on structural changes in the brain\u0026rsquo;s grey matter and in psychological aspects in two generations of people who were directly or indirectly affected by the war in the countries of the former Yugoslavia.\u003c/p\u003e\u003cp\u003eSurvivors of conflicts in the former Yugoslavia (G1 group)\u003c/p\u003e\u003cp\u003eIn this study, the most significant finding of the voxel-based morphometry analysis in G1 was a reduction in grey matter volume in the left cerebellar Crus II. Although the cerebellum has historically been associated primarily with motor functions \u003csup\u003e\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e\u003c/sup\u003e, recent research has confirmed its involvement in non-motor processes, including cognition, emotion, social cognition, and autobiographical memory \u003csup\u003e\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e,\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e\u003c/sup\u003e. The posterior lobe structure Crus II, part of the cerebellum\u0026rsquo;s phylogenetically newer regions, has been linked to social mentalisation, expression of self-related emotions, and the processing of autobiographical memory \u003csup\u003e\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e,\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e\u003c/sup\u003e. A recent meta-analysis identified a consistent association between PTSD and reduced volume in Crus II, with symptom severity showing a stronger correlation than diagnosis alone \u003csup\u003e\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e\u003c/sup\u003e. Functional MRI studies have also shown increased activation of Crus II during the recall of sad and negative autobiographical memories \u003csup\u003e\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e\u003c/sup\u003e. The posterior cerebellum is anatomically connected to paralimbic areas and contributes to the integration of emotion and behaviour \u003csup\u003e\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e\u003c/sup\u003e. A reduction in volume was also found in the right and left parahippocampus, part of the paralimbic system. The parahippocampal gyrus (PHG), located on the ventromedial surface of the temporal lobe and closely associated with the hippocampus, plays a role in emotional processing and memory consolidation \u003csup\u003e\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e\u003c/sup\u003e. Nardo and his team focused their study on grey matter concentration; they found a reduction in density in the PHG (and in the posterior cingulate and insular cortex) in PTSD subjects. They suggest that the altered function of PHG may lead to memory impairments seen in PTSD or impaired processing of emotional stimuli and integration of traumatic memories \u003csup\u003e\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e\u003c/sup\u003e. Also, reduction in the volume of this structure has been found in patients with PTSD \u003csup\u003e\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e\u003c/sup\u003e. In another work \u003csup\u003e\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e\u003c/sup\u003e, the authors argue that disrupted parahippocampal-prefrontal coupling may result in altered memory suppression in patients with PTSD \u003csup\u003e\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e\u003c/sup\u003e. A reduction in grey matter was also observed in the middle temporal gyrus (MTG), which is involved in cognition and memory processing, particularly in the retrieval and suppression of emotional memories, including PTSD flashbacks \u003csup\u003e\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e\u003c/sup\u003e. Volume reduction in the left MTG has been associated with impaired retrieval of autobiographical memories in PTSD \u003csup\u003e\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e\u003c/sup\u003e. The fusiform gyrus (FG), known for its role in face and body recognition and complex visual processing, is also involved in memory, emotional perception, and threat detection. Reduced volume in this region has been linked to deficits in visual and emotional processing of social stimuli in PTSD, potentially contributing to symptoms such as hypervigilance and social dysfunction \u003csup\u003e\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e,\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e,\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e\u003c/sup\u003e. Although the postcentral gyrus, which is primarily responsible for somatosensory processing, is not usually examined in research on psychological trauma, a study of war veterans with PTSD was conducted to examine somatosensory responses to non-threatening stimuli. Veterans with PTSD did not show typical responses to touch in the postcentral gyrus, superior parietal region, and right prefrontal cortex.Unlike participants with PTSD responded differently to non-threatening tactile stimuli This result suggests that PTSD manifests itself not only through psychological symptoms but also through somatosensory symptoms, such as altered responses to tactile stimuli \u003csup\u003e\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eThe finding that all regions in which we found volume reductions in G1 were associated with post-traumatic stress disorder or its symptoms corresponds with the results of the PCL-5 questionnaire, which revealed higher symptoms of persistent post-traumatic stress in G1. The threshold of 31 points or more \u003csup\u003e\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e\u003c/sup\u003e, on the basis of which a preliminary diagnosis of PTSD can be made, was exceeded in approximately half of the subjects in group G1. Therefore, while it cannot be conclusively stated that individuals in G1 meet the full diagnostic criteria for PTSD, they do exhibit pronounced symptoms indicative of post-traumatic stress. Furthermore, we found lower satisfaction with life (SWSL) in G1 as compared to CG. Similar results were found in an earlier study that examined individuals affected by the conflict in the former Yugoslavia 10 years later. These individuals still showed symptoms of post-traumatic stress decreased quality of life 10 years after the conflict \u003csup\u003e\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e\u003c/sup\u003e. Our results extend these findings to a broader time horizon. From our previous work examining Holocaust survivors \u003csup\u003e\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e\u003c/sup\u003e, we know that increased stress symptoms and lower well-being can persist over the long term and even throughout an entire lifespan. However, the same study found that post-traumatic growth also persists, as is the case with G1 in the current study. The coexistence of post-traumatic growth and stress is not an uncommon phenomenon. Some studies have explored the possibility that PTG is not a completed event based on an experienced event, but a process of seeking to cope with stress \u003csup\u003e\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eGiven that the reduction in grey matter volume in G1 occurred in the Crus II area, which is demonstrably involved in the processing of negative memories and social mentalisation, one plausible explanation for this reduction is the suppression of negative experiences, which may function as a coping mechanism in G1. This hypothesis is indirectly supported by interviews from our G2 participants, who are the children of war survivors, even though they are not direct descendants of our G1 participants. In interviews, G2 often reported that discussions of the war were common in their households. However, these conversations typically lacked deep introspection regarding the emotional experiences of their parents.\u003c/p\u003e\u003cp\u003eEmpirical studies suggest that while thought suppression or avoidance may offer short-term relief, it is associated with the persistence of PTSD symptoms in the long term \u003csup\u003e\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e\u003c/sup\u003e. This finding is consistent with the psychological questionnaires used in this study, which indicate that G1 participants exhibit higher levels of PTSD symptoms and lower life satisfaction. However, results from the PTGI suggest that, despite these challenges, G1 participants also experience post-traumatic growth, reporting positive evaluations of their personal and professional lives. Many have achieved above-average incomes, high levels of education, and have successfully integrated into countries other than their homeland.\u003c/p\u003e\u003cp\u003eAlthough suppression is generally considered a maladaptive long-term coping strategy, it does not preclude the possibility of resilience. In their research on resilience in post-war Southeastern Europe, Kelmendi and Hamby identified emotional self-regulation as a prominent trauma coping mechanism characteristic of this region \u003csup\u003e\u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e\u003c/sup\u003e. In G1, the coping mechanism appears to involve conscious or unconscious suppression of negative autobiographical memories, leading to the creation of a \u0026ldquo;protective shell\u0026rdquo; that allows them to cope with trauma and live a full life, without completely eliminating their emotional scars.\u003c/p\u003e\u003cp\u003eChildren of survivors (G2 group)\u003c/p\u003e\u003cp\u003eThe second generation in this study consisted of people who were born after the end of the war to parents affected by the war (in Bosnia and Herzegovina or in Croatia) or the bombing in Serbia in 1999. Most participants grew up in the post-war countries of the former Yugoslavia, some were born elsewhere, and a few participants were born and spent their childhood in the Czech Republic, although they continued to have a relationship with their parents\u0026rsquo; countries of origin. In the interviews, the participants reported that the topic of war is still alive. It was repeatedly mentioned that for some families, the war is a constant part of almost every conversation, and time is split into pre-war and post-war periods, with parents often talking about what happened in terms of their perspective of everyday life but sharing little about their inner experiences and feelings.\u003c/p\u003e\u003cp\u003eWe found no differences between the brain structures of the G2 and their peers in CG2. Neither were differences found in the PCL-5 and SWSL questionnaires: participants do not show differences in persistent symptoms of post-traumatic stress nor feel lower satisfaction with life compared to the control group. Like G1, G2 do show higher post-traumatic growth than CG. However, participants from G2 did not have a significant difference in PCL-5, nor did they exceed the cut-off threshold for PCL-5 in greater numbers (only three subjects had scores above 31). Thus, it cannot be said that this group carried persistent post-traumatic stress.\u003c/p\u003e\u003cp\u003eThe absence of post-traumatic stress and the presence of post-traumatic growth are not mutually exclusive. Although in most interviews, with the exception of two participants, participants in G2 rated the war as having had an impact on them, a possible explanation is that they have successfully integrated the difficulties they experienced into their lives, or that PTG has served as an adaptation process, as we indicated above for G1 \u003csup\u003e47\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eBased on our recent research, we found that the second generation differed in some ways from the non-war-affected control group, but we could not determine whether this was due to the post-war environment \u003csup\u003e\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e\u003c/sup\u003e, family and collective history \u003csup\u003e\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e51\u003c/span\u003e\u003c/sup\u003e, transgenerational transmission, or a combination of these factors. It seems that these questions will be the subject of new research in the future \u003csup\u003e\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eLimitations\u003c/p\u003e\u003cp\u003e\u003cul\u003e\u003cli\u003e\u003cp\u003eOur group cannot represent an independent cross-section of the Balkan population. In our research, we only worked with individuals who volunteered for the study; their motivation to participate itself may have biased the sample. Furthermore, these people have an above-average socioeconomic background; G1 participants had lived in a country other than their home country for a long time.\u003c/p\u003e\u003c/li\u003e\u003cli\u003e\u003cp\u003eG1 participants had experienced war in many forms and in different countries of the former Yugoslavia (Bosnia and Herzegovina, Croatia, Serbia). It is not a perfectly homogeneous group, but on the other hand, it is a model sample of people affected by war in various forms.\u003c/p\u003e\u003c/li\u003e\u003cli\u003e\u003cp\u003e Our two stress groups, G1 and G2, are not parents and children. Therefore, we cannot observe direct transmission from parents to offspring.\u003c/p\u003e\u003c/li\u003e\u003c/ul\u003e\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eIndividuals affected by the war in the former Yugoslavia exhibit reduced grey matter volume in brain structures associated with post-traumatic stress disorder (PTSD) symptoms and autobiographical memory processing. These individuals continue to experience persistent symptoms of post-traumatic stress and report lower life satisfaction compared to control participants. At the same time, they demonstrate higher levels of post-traumatic growth and, in self-reports, assess their lives positively. This pattern suggests the presence of a specific, albeit imperfect, coping mechanism that enables a more reactive response to perceived or actual threats, while still allowing for the pursuit of a fulfilling life. In contrast, the second generation, raised in a challenging post-war environment shaped by a pervasive war narrative, do not exhibit structural brain changes or significant differences in life satisfaction and stress symptoms compared to their peers in the control group. Although they do not show persistent symptoms of post-traumatic stress, they tend to perceive that the difficulties caused by the post-war environment have a negative impact on their lives. At the same time, they report increased post-traumatic growth, suggesting that they may have effectively integrated these challenges into their lives or that post-traumatic growth serves as an adaptive ongoing mechanism supporting their ability to cope with adversity.\u003c/p\u003e\u003cp\u003eFinally, these findings highlight the need for stress research to consider the role of the cerebellum, particularly Crus II, which has long been overlooked in studies of non-motor brain functions\u003c/p\u003e"},{"header":"Methods and analysis","content":"\u003cp\u003e\u003cem\u003eResearch\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe research was conducted between 2022 and 2025 at the Central European Institute of Technology—Research Centre of Masaryk University in Brno. It was approved by the Ethics Committee of Masaryk University in accordance with the Declaration of Helsinki. The ethics committee’s approval code number is EKV-2021-076. Informed consent was obtained from each participant.\u003c/p\u003e\n\u003cp\u003eThis project represents an interdisciplinary research effort examining the psychological, physiological, and neuroimaging correlates of war-related stress in civilian populations. The current study specifically focuses on data pertinent to the assessment of brain structure, utilising structural MRI findings alongside psychological questionnaires and interview data.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eRecruitment\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eVolunteers were recruited between 2022 and 2024 through media reports, university information channels, social networks, and a lecture at the Lastavica club, which brings together the Balkan diaspora in the Czech Republic.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eParticipants\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWe investigated participants who, as civilians, survived the war in former Yugoslavia and second-generation survivors – children of war survivors. The group of participants we categorised as the first generation (G1) was composed of people (n = 45) who experienced multiple war events and were under life-threatening stress during war activity in then-Yugoslavia in 1991–1995 (Bosna and Herzegovina, Croatia) or experienced NATO bombing in Serbia in 1999, or, in some cases, a combination of these events. To the question, \u003cem\u003eHow did you survive the war? What did you experience during the war?\u003c/em\u003e participants provided a variety of responses, including:\u003c/p\u003e\n\u003cul\u003e\n \u003cli\u003e\n \u003cp\u003eShelling and bombing - Hiding in a cellar, shelter, or other hiding places\u003c/p\u003e\n \u003c/li\u003e\n \u003cli\u003e\n \u003cp\u003eForced abandonment or destruction of homes\u003c/p\u003e\n \u003c/li\u003e\n \u003cli\u003e\n \u003cp\u003eWar-related injuries - Death or injury of loved ones\u003c/p\u003e\n \u003c/li\u003e\n \u003cli\u003e\n \u003cp\u003eWitnessing the death or injury of others\u003c/p\u003e\n \u003c/li\u003e\n \u003cli\u003e\n \u003cp\u003eSeparation from family and loved ones\u003c/p\u003e\n \u003c/li\u003e\n \u003cli\u003e\n \u003cp\u003eFear for the safety of loved ones\u003c/p\u003e\n \u003c/li\u003e\n \u003cli\u003e\n \u003cp\u003eLack of food, medicine, water, and electricity\u003c/p\u003e\n \u003c/li\u003e\n \u003cli\u003e\n \u003cp\u003ePoverty\u003c/p\u003e\n \u003c/li\u003e\n \u003cli\u003e\n \u003cp\u003eDiscriminatory and humiliating behaviour by soldiers\u003c/p\u003e\n \u003c/li\u003e\n \u003cli\u003e\n \u003cp\u003eInvoluntary service in the military (none of the participants were professional soldiers)\u003c/p\u003e\n \u003c/li\u003e\n \u003cli\u003e\n \u003cp\u003eInconveniences associated with forced emigration, including bullying or rejection, loss of social capital, and financial difficulties.\u003c/p\u003e\n \u003c/li\u003e\n\u003c/ul\u003e\n\u003cp\u003eAfter the experience of the conflict, these people emigrated from their home country directly to the Czech Republic or to other countries and then to the Czech Republic, where they currently live.\u003c/p\u003e\n\u003cp\u003eThe second generation (G2) contains people who are descendants of war survivors (n = 28) and were not exposed to war-threatening stress but grew up in a post-war environment. In addition to five persons of Czech nationality who were born in the Czech Republic to parents who had emigrated to the Czech Republic, there were participants of Bosnian, Croatian, Serbian and Montenegrin nationality who were temporarily in the Czech Republic as part of a study programme. G1 and G2 are not family members (with a few exceptions). G2 participants are not direct descendants of G1.\u003c/p\u003e\n\u003cp\u003eThe control group (CG) consisted of Czechs or Slovaks living in Czechia who had no war experienc; the group were divided into two subgroups corresponding to G1 and G2, we called the subgroups CG1 and CG2 (n = 73; 45 + 28).\u003c/p\u003e\n\u003cp\u003eExclusion criteria\u003c/p\u003e\n\u003cp\u003eBrain impairment (brain injuries or diseases, tumours, neurodegenerative diseases), severe psychiatric disorders*(e.g. psychosis), significant cognitive decline. Furthermore, participants with technical artefacts or excessive movement artefacts in the MR images were excluded.\u003c/p\u003e\n\u003cp\u003e* Due to the nature of the research, individuals who were previously diagnosed with post-traumatic stress disorder or believed they might be diagnosed with it were retained in the sample.\u003c/p\u003e\n\u003cp\u003eThe stress groups and their assigned control groups did not differ significantly in age, sex, or education. Income was significantly higher in G1 than in CG1. The education and income of G2 and CG2 were not assessed because most participants were students. Age was compared using the two-sample t-test; sex, education, and income were compared using the Fisher exact test (Table\u0026nbsp;4).\u003c/p\u003e\n\u003cdiv\u003e\n \u003ctable id=\"Tab4\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv\u003eTable 4\u003c/div\u003e\n \u003cdiv\u003e\n \u003cp\u003eDemographic data of the research population\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\u0026nbsp;\u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eG1\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCG1\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ep-value\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eCount\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e45\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e45\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eAge\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e38 ± 9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e37 ± 10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.5870\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eSex\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23 M / 22 F\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20 M / 25 F\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.6732\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eUniversity education\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e29 Y / 16 N\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e32 Y / 13 F\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.6523\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eIncome\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e30 H / 15 L\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e11 H / 34 L\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003e0.0001\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eG2\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eCG2\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003ep-value\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eCount\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e28\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e28\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eAge\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e25 ± 3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e24 ± 3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.3255\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eSex\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e9 M / 19 F\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e11 M / 18 F\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.7828\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003e\u003cem\u003eMRI\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eMR examinations were performed on a 3T scanner Siemens Prisma using a 64-channel head coil. The MRI protocol for voxel-based morphometry included 3D T1-weighted magnetisation prepared rapid gradient echo (MPRAGE) sequence with TR = 2.3 s, TE = 2.33 ms, TI = 0.9 s, FA = 8°, isometric voxel size 1 mm in FOV 224 × 224 mm and 240 slices.\u003c/p\u003e\n\u003cp\u003eAnatomical MRI data were analysed using SPM12 (www.fil.ion.ucl.ac.uk) and CAT12 toolbox (www.neuro.uni-jena.de/cat) running in Matlab R2020a.\u003c/p\u003e\n\u003cp\u003eThe standard recommended pipeline was used for segmentation, in short: high resolution data were segmented into grey matter (GM) using the SPM Tissue Probability Map (TPM) and registered into common MNI space using shooting template IXI555_MNI152_GS. Finally, spatially normalised and modulated GM maps were smoothed with 6 mm FWHM isotropic Gaussian kernel.\u003c/p\u003e\n\u003cp\u003eGroup-level statistics for stress effects were computed using a second-level model in SPM12. Modulated GM images were corrected for total intracranial volume and subsequently analysed. A two-sample t-test compared GMV maps between the stress groups (G1, G2) and control groups (CG1, CG2); sex and age were included as nuisance variables.\u003c/p\u003e\n\u003cp\u003eResultant t-statistic maps were initially thresholded at a P value of \u0026lt; 0.001 uncorrected and then only significant clusters at P \u0026lt; 0.05 FWE cluster level were picked.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003ePsychological questionnaires \u0026amp; Self-report \u0026amp; Interview\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eParticipants completed a set of psychological questionnaires (Table\u0026nbsp;5), a self-assessment, and a semi-structured interview, allowing them to answer questions briefly or provide more detailed responses if they wished.\u003c/p\u003e\n\u003cdiv\u003e\n \u003ctable id=\"Tab5\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv\u003eTable 5\u003c/div\u003e\n \u003cdiv\u003e\n \u003cp\u003ePsychological questionnaires\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePTGI\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePost-Traumatic Growth Inventory\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eAssesses positive outcomes of experiencing stressful life events. The Inventory contains 21 items and measures 5 factors: New Possibilities, Relating to Others, Personal Strength, Spiritual Change, and Appreciation of Life. Participants were asked to mark the degree of perceived change (0 = I did not experience this change as a result of my crisis to 5 = I experienced this change to a very great degree). The score range is from 0 to 105, with higher scores indicative of greater post-traumatic growth \u003csup\u003e53\u003c/sup\u003e.\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePCL-5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eThe PTSD Check List for DSM-5\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eA 20-item self-report instrument for measuring symptoms of PTSD based on DSM-5. Respondents are asked to consider a “list of problems and complaints that people sometimes have in response to stressful experiences” that is supposed to rate how much they “have been bothered by each problem in the past month”. Every item is scaled from 0 (“not at all”) to 4 (“extremely”). Completing the inventory takes approximately 5–10 minutes. The PLC-5 questionnaire can be evaluated in several ways. A cut-off score between 31 and 33 items is used for a provisional diagnosis of PTSD \u003csup\u003e45\u003c/sup\u003e.\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSWSL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eSatisfaction with Life Scale\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eA 5-item self-report instrument. Respondent answers are marked on Likert scale ranging from 1 (“strongly disagree”) to 7 (“strongly agree”), and are presented in raw scores, with a total score range from 5 to 35. The SWLS focuses on cognitive aspects of life satisfaction rather than emotional; moreover, SWLS does not focus on specific domains. SWLS makes it possible to weigh different aspects of life based on personal criteria \u003csup\u003e54\u003c/sup\u003e.\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eMSPSS\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eMultidimensional Scale of Perceived Social Support\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eA 12-item self-report scale. The instrument measures subjectively assessed social support based on three subscales addressing a different source of support: a) family, b) friends and c) significant others. Respondent answers are marked on Likert scale ranging from 1 (“very strongly disagree”) to 7 (“very strongly agree”) \u003csup\u003e55\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cstrong\u003eBrief COPE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eA 28-item self-report inventory focusing on effective and ineffective ways to cope with stressful events. It is useful tool for detecting emotional responses to serious events e.g. health conditions, financial stress, mental illness, injuries and more. Respondent answers were marked on scale ranging from 1 (“I haven't been doing this at all”) to 4 (“I've been doing this a lot”) \u003csup\u003e56\u003c/sup\u003e.\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003eThe results of the psychological questionnaires were summarised using the median, as well as the Interquartile range (IQR). To test for statistical differences, the non-parametric Mann-Whitney U test was employed. The significance level for all statistical tests was set at p \u0026lt; 0.05.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSelf-report \u0026amp; interview\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll participants completed a short self-assessment of their life and work achievements to date. Satisfaction with life and career was rated on a scale of yes, rather yes, somewhere in between, rather no, no. Participants from G2 did not rate career success as they were predominantly students. Stress groups (G1 and G2) additionally provided information on war experiences affecting themselves or their families through semi-structured interviews, forming the basis for a separate psychological study. For the current text, we report only demographic data, life satisfaction ratings, and war-related experiences. G1 participants were asked whether war was the worst event of their life, and how they coped; G2 participants were asked whether they felt that the war had affected them indirectly and whether the war had affected their parents’ parenting style.\u003c/p\u003e"},{"header":"Abbreviations","content":"\u003cp\u003eG1 - First Generation\u003c/p\u003e\n\u003cp\u003eG2 - Second Generation\u003c/p\u003e\n\u003cp\u003eCG - Control Group\u003c/p\u003e\n\u003cp\u003eCG1 - Control Group for G1\u003c/p\u003e\n\u003cp\u003eCG2 - Control Group for G2\u003c/p\u003e\n\u003cp\u003eMRI - Magnetic Resonance Imaging\u003c/p\u003e\n\u003cp\u003ePTSD - Post-Traumatic Stress Disorder\u003c/p\u003e\n\u003cp\u003ecPTSD \u0026nbsp;- Complex Post-Traumatic Stress Disorder\u003c/p\u003e\n\u003cp\u003eICD-11 - International Classification of Diseases, 11th Revision\u003c/p\u003e\n\u003cp\u003ePTGI - Post-Traumatic Growth Inventory\u003c/p\u003e\n\u003cp\u003ePCL-5 - PTSD Checklist for DSM-5\u003c/p\u003e\n\u003cp\u003eSWSL - Satisfaction with Life Scale\u003c/p\u003e\n\u003cp\u003eMSPSS - Multidimensional Scale of Perceived Social Support\u003c/p\u003e\n\u003cp\u003eBrief-COPE - Brief Coping Orientation to Problems Experienced Inventory\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis study was supported by a grant from the Ministry of Health of the Czech Republic, grant no. NU22-04-00661.\u003c/p\u003e\n\u003cp\u003eData availability\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe datasets used and/or analysed during the current study available from the corresponding author on reasonable request. The authors assert that all procedures contributing to this work comply with the ethical standards of the relevant national and institutional committees on human experimentation and with the Helsinki Declaration of 1975, as revised in 2008.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAcknowledgement\u003c/p\u003e\n\u003cp\u003eWe thank Anne Johnson for grammatical assistance. We acknowledge the core facility MAFIL of CEITEC MU.\u003c/p\u003e\n\u003cp\u003eWe acknowledge the core facility MAFIL supported by the Czech-BioImaging large RI project (LM2023050 funded by MEYS CR), part of the Euro-BioImaging (www.eurobioimaging.eu) ALM and Multimodal Imaging Node (Brno, CZ), for their support with obtaining scientific data presented in this paper.\u003c/p\u003e\n\u003cp\u003eAuthor information\u003c/p\u003e\n\u003cp\u003eAuthors and Affiliations\u003c/p\u003e\n\u003cp\u003eCentral European Institute of Technology (CEITEC), Brain and Mind Research Programme, Masaryk University, Brno, Czechia\u003c/p\u003e\n\u003cp\u003eMonika Fňa\u0026scaron;kov\u0026aacute;, Pavel Ř\u0026iacute;ha, Marek Preiss, David Ulč\u0026aacute;k, Mark\u0026eacute;ta Nečasov\u0026aacute;, Nikola Wolframov\u0026aacute;, Martin Lamo\u0026scaron; \u0026nbsp;\u0026amp; Ivan Rektor\u003c/p\u003e\n\u003cp\u003eNational Institute of Mental Health (Czechia), Prague, Czechia\u003c/p\u003e\n\u003cp\u003eMarek Preiss\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;3. \u0026nbsp;First Department of Neurology, St Anne\u0026rsquo;s University Hospital and Faculty of \u0026nbsp; Medicine, Masaryk University, Brno, Czechia\u003c/p\u003e\n\u003cp\u003eMonika Fňa\u0026scaron;kov\u0026aacute;, Pavel Ř\u0026iacute;ha, David Ulč\u0026aacute;k, Martin Lamo\u0026scaron; \u0026nbsp;\u0026amp; Ivan Rektor\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;4. University of New York in Prague\u003c/p\u003e\n\u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; Marek Preis\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;4. Utrecht University\u003c/p\u003e\n\u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; Vojtěch Svoboda\u003c/p\u003e\n\u003cp\u003eContributions\u003c/p\u003e\n\u003cp\u003eM.F. (corresponding author): Project Administration, Investigation, Resources, Data Curation, Writing Original Draft, Visualisation, P.Ř. - Methodology, Software, Formal Analysis, Investigation, Visualisation, M.P. - Conceptualisation, Methodology, Validation, D. U. Investigation, Visualisation, M.N. - Investigation, N.W. Investigation, V. S. Investigation, M.L. Supervision, I.R. - Conceptualisation, Supervision.\u003c/p\u003e\n\u003cp\u003eCorresponding author\u003c/p\u003e\n\u003cp\u003eCorrespondence to Monika Fňa\u0026scaron;kov\u0026aacute; ([email protected])\u003c/p\u003e\n\u003cp\u003eEthics declarations\u003c/p\u003e\n\u003cp\u003eCompeting interests\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eEthical approval for this study was obtained from the ethics committee of Masaryk University (approval code EKV-2021-076) on June 24, 2021\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eComtesse, H., Powell, S., Soldo, A., Hagl, M. \u0026amp; Rosner, R. 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This study focuses on survivors of the war in the former Yugoslavia (G1) who now reside in the Czech Republic and on the children of survivors (G2) who were born after the conflict.\u003c/p\u003e\u003cp\u003e Participants from G1, G2, and a control group (CG) with no war experience underwent structural MRI, answered a semi-structured interview, and completed psychological questionnaires (PTGI, PCL-5, SWSL, MSPSS, Brief-COPE). Voxel-based morphometry was used to assess brain volume differences.\u003c/p\u003e\u003cp\u003eCompared to CG, G1 showed reduced grey matter volume in regions associated with PTSD and autobiographical memory, including Crus II, parahippocampal gyrus, middle temporal gyrus, and fusiform gyrus. Psychologically, G1 reported higher PTSD symptoms, lower life satisfaction, and greater post-traumatic growth. G2 showed no structural brain changes but scored higher on post-traumatic growth than CG, with no significant differences in other psychological measures.\u003c/p\u003e\u003cp\u003eThe findings suggest long-lasting neuroanatomical and psychological effects of war stress in directly exposed individuals (G1). Although G2 showed no brain alterations, the increased post-traumatic growth may indicate subtle adaptation effects of growing up in a post-war environment, but not necessarily only in a maladaptive sense.\u003c/p\u003e","manuscriptTitle":"Long-term impact of war stress in civilians from the former Yugoslavia: A two-generation neurostructural and psychological study","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-08-05 06:53:53","doi":"10.21203/rs.3.rs-7028508/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-09-15T19:26:23+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-08-25T14:08:52+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-08-15T20:28:21+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"76701529894148498283453751643321827973","date":"2025-08-15T00:38:14+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"302412200490488819769857202686723476488","date":"2025-08-06T23:46:56+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-07-30T16:32:24+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-07-22T16:02:40+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2025-07-07T17:43:29+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-07-07T09:27:55+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2025-07-02T10:38:38+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"172f43dc-7387-40ed-8e36-976accd96bdc","owner":[],"postedDate":"August 5th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":52563233,"name":"Biological sciences/Neuroscience"},{"id":52563234,"name":"Biological sciences/Psychology"},{"id":52563235,"name":"Social science/Psychology"}],"tags":[],"updatedAt":"2026-03-23T16:03:33+00:00","versionOfRecord":{"articleIdentity":"rs-7028508","link":"https://doi.org/10.1038/s41598-026-44241-w","journal":{"identity":"scientific-reports","isVorOnly":false,"title":"Scientific Reports"},"publishedOn":"2026-03-17 15:59:43","publishedOnDateReadable":"March 17th, 2026"},"versionCreatedAt":"2025-08-05 06:53:53","video":"","vorDoi":"10.1038/s41598-026-44241-w","vorDoiUrl":"https://doi.org/10.1038/s41598-026-44241-w","workflowStages":[]},"version":"v1","identity":"rs-7028508","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7028508","identity":"rs-7028508","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}