Investigation and validation of a natural depression model in cynomolgus monkeys （Macaca fascicularis）

Investigation and validation of a natural depression model in cynomolgus monkeys （Macaca fascicularis） | 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 Research Article Investigation and validation of a natural depression model in cynomolgus monkeys （Macaca fascicularis） Wei Li, Li Huang, Shi-Gang Liu, Jun Tang This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6716220/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 16 Oct, 2025 Read the published version in BMC Zoology → Version 1 posted 11 You are reading this latest preprint version Abstract Background The global burden of depression is heavy, with unclear pathogenesis, relying on animal models for researching neural mechanisms and treatments. Existing rodent models lack social attributes, while non-human primate models (such as the natural depression model in cynomolgus monkeys) are closer to human social stress scenarios. Previously, the screening of natural depression models in cynomolgus monkeys was unvalidated and their causes were unclear. This study constructs a cynomolgus monkey community by simulating wild environments, records social behaviors, and explores the causes of their natural depression-like behaviors and validates them, providing theoretical support for this model. Results The conflict behaviors in the community were recorded to develop a conflict behaviors matrix, and the behaviors of all individuals were monitored by using the focus observation method. The analysis entailed recording and analyzing ten behaviors for ten observation time slots on individuals with depression. The David Scoring (DS) scores of the animals were determined based on the record of conflict behaviors in the community. The individuals were divided into high- and low-level groups, and the daily behaviors of the two groups were analyzed. The results showed statistically significant differences in depression behaviors, anxious behaviors, and motor behaviors among the monkeys. Further, individuals with depression behaviors exhibited drastically lower capability in exploration and social interaction. Additionally, social frustration stress responses induced by inter- and intra-community conflict behaviors may account for this behavior in the cynomolgus monkey model of natural depression. Conclusions The natural depression model of caged Macaca fascicularis is a social defeat model induced by yield-attack behavior. MDD Macaca fascicularis Chronic unpredictable mild stress Social interaction test Figures Figure 1 Figure 2 Background According to 2019 statistics released by the World Health Organization (WHO), 320 million depression cases were reported globally by 2019, of which 95 million depression cases were reported in China. Furthermore, research has shown that depression ranks second among the largest disease burdens in China [ 1 ]. There are no clear experimental indicators because of the inherent limitations of human studies and the unclear pathogenesis of depression. Hence, the use of animal models remains crucial in identifying the neural mechanisms of depression and discovering new drugs and treatments. Currently, rodent animal models, such as the chronic unpredictable mild stimulus model, behavioral despair model, and the social frustration stress model, are used in basic depression research [ 2 ]. However, these animal models are established in non-social environments. However, depression is a disease closely related to social pressure and social relationships. Further, rodent animal models have several limitations, including the lack of complex characteristics seen in human society. Non-human primate models of depression can compensate for this challenge of rodent systems. Currently, non-human primate models of depression include the social isolation model, mother-infant separation model, and natural depression model [ 3 ]. Among these models, the natural depression model is the most ethical and socially relevant one. The approach identifies animals with eye-opening, crouching, and head bowing as core behaviors in a caged community of cynomolgus monkeys (Macaca fascicularis) [ 4 ]. For instance, Shively et al. confirmed a few monkeys with natural depression, although the causes of natural depression in the animals were not specified [ 5 – 6 ]. Further, even when this natural depression model was screened, it was not validated. Given these findings, a community of cynomolgus monkeys was established by mimicking wild conditions. The social behaviors of cynomolgus monkeys were recorded, and the causes of their depression-like behaviors were investigated. Further, the interactive behavior and food exploration behavior were evaluated to validate the results. This study aims to examine the causes of natural depression behaviors and validate them with corresponding evaluations. The results can provide reliable theoretical support for future studies of cynomolgus monkey models of natural depression. Materials and methods Animal details and selection This study was conducted at the non-human primates laboratory of Suzhou Zhongke Animal Experiment Co., Ltd, certified by the Association For Assessment and Accreditation of Laboratory Animal Care (AAALAC). The cynomolgus monkeys were provided by the Zhongke Animal Experimental Base. The license number for the domestication and breeding of nationally protected wild animals was SuFa Domestication 2007-001. Initially, 22 healthy cynomolgus monkeys from different cages were randomly selected from the Zhongke Animal Experimental Base. After excluding pregnancy and other interfering factors, 19 cynomolgus monkeys were selected to form a community. The average age was 8.4 ± 1.1 years, and the male-to-female ratio was 1:8.5. The housing met the cage standards for caged cynomolgus monkeys in China. The basic data of the cynomolgus monkeys in this study is shown in Table 1 . Table 1 The basic data of the cynomolgus monkeys( Macaca fascicularis ) Number Experiment No. Gender Age (years) Number Experiment No. Gender Age (years) 1 12152 F 7 11 06121 F 7 2 11192 F 8 12 05117 F 8 3 08264 F 8 13 01102 F 7 4 03150 F 10 14 11240 F 7 5 12052 F 9 15 03312 F 9 6 04126 F 9 16 07258 F 9 7 05056 F 9 17 05128 F 9 8 01140 F 6 18 07162 M 10 9 03128 F 9 19 12118 M 7 10 08198 F 8 Methods Establishment of the cynomolgus monkey community Cynomolgus monkeys may be exposed to fierce conflict behaviors due to competition for food resources, territory, and mating rights [ 7 – 8 ]. In this study, a community of “multiple females and multiple males” was created by simulating the wild living habitats of cynomolgus monkeys. Observing the daily behaviors and conflict behaviors The characteristics of cynomolgus monkey individuals were identified in the pre-observation phase conducted for 2 weeks to provide an accurate basis for the subsequent recording of various behaviors. After the pre-observation period, each community of cynomolgus monkeys was assigned four experimental recorders; two were observed for attack-yield behaviors, and the other two were photographed and observed for daily behaviors. Different behaviors were observed in ten time slots, and a consistency test was performed among the behaviors, with k > 80% as valid observations. The observation researchers recorded the effective number of attack-yield behaviors according to the behavioral recording items. The researchers who observed the daily behavior followed and photographed cynomolgus monkeys using the focused filming method [ 9 ] .The observation time was 8:30 − 11:30 and 14:00–16:00; each time slot was 30 min, with ten time slots per day, and the cumulative effective time was 1050 h. The Noldus analysis system was used to analyze animal behavior [ 10 ]. Record of conflict behaviors The conflict behaviors are defined in Table 2 based on the behavioral spectrum of cynomolgus monkeys [ 11 ]. Table 2 Definition of cynomolgus monkeys( Macaca fascicularis ) Conflict Behavior Behavior category Behavioral definition Attack behaviour Bite Aggressive behaviors such as biting each other's head, back, limbs, and other body parts with teeth represent the most severe form of aggression, which can result in fatal consequences for the attacked individual Grab, hit, push Minimal physical contact, infrequently resulting in bodily harm Threats Engaging in verbal altercations and forcefully shaking doors and windows with upper limbs Yielding behavior Run away Quickly flee from the attacker after being attacked and scream. Passive submissio Following an attack, the individual does not flee from the aggressor, instead emitting a scream and displaying friendly behaviors towards the attacker, such as grooming Avoidance The two parties do not engage in physical confrontation and deliberately distance themselves from their original positions Definition and records of daily behaviors The animal behavior analysis system was used to record and analyze ten types of behaviors, including anxiety, conflicts, and food intake [ 12 ]. Friendly behaviors refer to individuals grooming other individuals after an active attack or being attacked. Anxious behaviors are noted in individuals who rapidly scratch for more than 3 s. Rest behaviors refer to standing, sitting, or lying down for more than 5 s. Conflict behaviors include biting, scratching, or threatening another individual. Food intake behaviors refer to individuals grabbing food from the trough. Warming behaviors refer to individuals snuggling with each other to maintain body temperature. Mating behaviors refer to sexual behaviors between individuals of opposite sexes. Motor behaviors include moving quickly or slowly, with a distance greater than half an arm’s length. Depression behaviors refer to the presence of core depression-like behaviors, including eye-opening, crouching, and head- bowing. Finally, other behaviors refer to behaviors other than those in the above nine categories. Food exploration test Individuals were divided into high- and low-level groups based on their David Scoring (DS) scores. Food (apples) was placed in the trough of a separate cage (2m ×1m ×1m) and buried with stones. A high-definition camera was placed 1 m in front of the cage to record the behavior of the cynomolgus monkeys with the apples without any interference. After turning on the camera, the experimenters left the cage, and recorded the time from placing the food to the time when the subject touched the food for each monkey. The subject animal was familiarized with the cage environment before the experiment, as shown in Fig. 1 . Interactive behavioral evaluation Single cages (2m×1m×1m) were separated from each other by a transparent plastic partition with holes of 1 cm in diameter to avoid contact between subjects but not to isolate them from sight and smell. Further, high- and low-level groups from each community were tested separately. The interaction subject was a healthy male cynomolgus monkey from another community, and the effective contact area was defined as the area 0.5 m to the left and right of the transparent partition. Each experiment lasted 5 min, and the time spent in the effective contact area by each cynomolgus monkey individual within 5 min from placing the male cynomolgus monkeys was recorded, as shown in Fig. 2 . Statistical analysis Data on the daily behavior indicators were expressed as means ± SD , and the Shapiro-Wilk test was used to test the normal distribution of the eleven behavioral aspects. As the data were not normally distributed, the nonparametric Wilcoxon rank sum test was used for the statistical analysis, and the significance level was set at α = 0.05. The attack-yield indicators were expressed as times, and the DS scores of each subject were calculated based on the attack-yield relationship matrix using the David’s score method. The data of food exploration tests were expressed as means ± SD and an independent samples t-test was conducted, with the significance level atα = 0.05. Results Attack-yield (conflict) behavior correlation matrix and David Scoring scores The established communities had significantly intense conflict behaviors since their formation, and 1122 valid conflict behaviors were recorded. David Scoring (DS) method was used to count the DS scores of each subject, and the formula was DS = W1 + W2 - L1 - L2. Here, W1 was the sum of the dominant percentages of monkeys who exhibited conflict behaviors in the group; W2 was the sum of the dominant weighted percentage in individual conflict behaviors; L1 was the sum of the disadvantage percentages of animals who exhibited conflict behaviors in the group; and L2 was the sum of the weighted disadvantage percentages in individual conflict behaviors [ 13 – 15 ]. The DS scores are shown in Table 3 . Table 3 Matrix of aggressive behavior relationships and individual DS scores for the group Attacker DS scores Genuflector M 1 M 2 F 1 F 2 F 3 F 4 F 5 F 6 F 7 F 8 F 9 F 10 F 11 F 12 F 13 F 14 F 15 F 16 F 17 M 1 M 2 F 1 F 2 F 3 F 4 F 5 F 6 F 7 F 8 F 9 F 10 F 11 F 12 F 13 F 14 F 15 F 16 F 17 - 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 171 6 - 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 152 12 13 - 1 0 0 0 0 0 0 0 0 3 0 0 0 0 0 0 131 7 22 8 - 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 116 14 5 8 5 - 0 0 0 0 0 0 0 0 0 3 0 0 0 0 95 11 2 4 7 3 - 0 0 0 0 0 0 0 0 0 0 0 0 0 73 11 5 7 19 3 9 - 0 0 0 0 0 0 0 0 0 0 0 0 57 15 3 11 12 2 5 7 - 0 0 0 0 0 0 0 3 0 0 0 4 16 6 7 9 3 6 6 12 - 0 0 0 0 0 0 0 0 2 0 19 11 6 3 6 2 10 2 29 2 - 0 0 0 0 0 0 0 0 0 1 10 3 12 4 2 2 2 22 8 2 - 0 0 0 0 0 0 0 0 -18 13 9 7 22 3 2 5 19 5 3 2 - 0 1 0 0 0 0 0 -38 8 3 9 8 2 11 2 17 10 2 5 2 - 0 0 0 0 0 0 -57 14 14 8 8 3 13 6 19 5 5 17 2 4 - 0 0 0 0 0 -88 9 2 3 4 4 15 3 8 4 3 16 3 3 3 - 0 0 0 0 -94 6 10 3 4 2 2 5 17 8 2 10 2 3 2 17 - 0 0 0 -119 4 4 3 5 3 4 2 3 5 3 7 3 6 3 2 3 - 0 0 -120 16 7 3 5 2 3 2 18 2 3 7 2 3 3 7 7 6 - 0 -150 5 6 3 11 2 8 3 6 3 6 14 2 2 2 13 6 22 5 - -171 Daily behaviors of attack and yield groups Individuals with positive and negative DS scores were divided into the high-level group (the attack group) and the low-level group (the yield group), respectively, and their daily behaviors were observed. The animals showed statistically significant differences in friendly, anxious, conflicting, mating, motor, and depression behaviors (P < 0.05), as shown in Table 4 . Table 4 Daily Observations of the Aggressor Group and the Yielding Group Behavioral items Aggressor Group (n = 10) Yielding Group (n = 9) t P Depression behaviors 1.01 ± 1.15 30.44 ± 23.90 -3.691 0.006* Friendly behaviors 49.30 ± 24.29 140.77 ± 48.79 -5.085 0.024* Mating behaviors 4.88 ± 1.54 1.88 ± 1.05 4.731 0.040* Conflict behaviors 76.30 ± 22.24 12.55 ± 7.97 8.478 0.018* Rest behaviors 800.60 ± 81.24 709.00 ± 50.92 0.663 0.518 Warming behaviors 23.50 ± 6.02 22.44 ± 5.27 0.503 0.551 Anxious behaviors 11.20 ± 8.23 73.11 ± 73.11 -4.053 0.005* Motor behaviors 141.1 ± 25.12 31.77 ± 24.17 9.639 0.007* Food intake behaviors 48.60 ± 14.55 7.88 ± 6.27 -0.284 0.780 Other behaviors 47.00 ± 12.69 50.11 ± 22.07 -0.382 0.707 Data are means ± SD . * : P < 0.05. Food exploration tests The time taken by the attack and yield groups to reach the food after placing apples was recorded. The results were statistically significant (P < 0.05), suggesting that low-level (yield) cynomolgus monkeys spent more time exploring the food than high-level (attack) ones (Table 5 ). Table 5 Comparison of the time taken by individuals of varying hierarchical positions within a group to explore food Aggressor Group Yielding Group t P n 10 9 Time taken to search for and find food 5.10 ± 2.12 15.03 ± 4.0 -0.422 0.015* Data are means ± SD . * : P < 0.05. The social interaction test in cynomolgus monkeys The time spent in the effective contact area within 5 minutes was recorded in high-level and low-level individuals in the group. The low-level (yield) individuals spent a significantly smaller time in the effective contact area than that of high-level (attack) monkeys, with statistically significant differences (P < 0.05) (Table 6 ). Table 6 Comparison of the time that individuals of various ranks within a group spend in the effective contact area Aggressor Group Yielding Group t P n 10 9 Time spent within the effective contact area 4.3 ± 1.07 1.01 ± 0.81 2.242 0.007 * Data are means ± SD . * : P < 0.05. Discussion Animal models with depression have always been vital for depression research. Further, the pathogenesis of depression can be better investigated with their high similarity with depression symptoms [ 16 ]. Currently, most animal models for depression use rodents, which lack certain social relevance. This lack can be compensated using non-human primate models of depression for this more accurate studies. For instance, Shively reported that some individuals spontaneously exhibited human-like depression due to social pressure in cynomolgus monkey communities [ 17 ]. However, the depression behaviors of cynomolgus monkeys are mainly reflected as crouching, eye-opening, and decreased mobility, which are different from those of humans. The cynomolgus monkey communities have strict social hierarchies, and the yield group is under long-term social pressure due to the lack of access to mating rights and food [ 18 ]. It is therefore hypothesized that these individuals are readily exposed to depression-like behaviors. In this study, we observed the attack-yield behaviors of cynomolgus monkeys after establishing a community. We recorded and analyzed the daily behaviors of individuals to explore the occurrence and the possible causes of depression-like behaviors in caged cynomolgus monkeys. In this study, 1122 behaviors were observed, and more intense attack-yield behaviors occurred from the beginning of group formation. Individuals were classified into the attack group and the yield group based on their DS scores, with significant differences in their daily behaviors. There were significant differences in the friendly, anxious, conflicting, mating, motor, and depression behaviors. The attack group was more likely to initiate conflicts, mating behavior, and motor behaviors with other individuals In contrast, the yield group was more likely to exhibit friendly, depressed, and anxious behaviors. The differences in group behaviors revealed the following characteristics: firstly, the attack group performed more prominently in attack behaviors than the yield group. The group attack-yield behavior correlation matrix showed that some community individuals could mitigate pressure by attacking and threatening other individuals when subjected to attacks and threats. In contrast, individuals with low DS scores had difficulty handling this pressure and were readily exposed to anxious and depressive behaviors. Further, the yield group was threatened by more individuals and rarely received friendly grooming behaviors from other individuals. The cynomolgus monkeys of the yield group developed more friendly behaviors to promote harmony with other individuals and avoid being attacked by others. Thirdly, as expected, the yield group developed typical solitary sitting, head bowing, crouching, and eating in more remote areas away from the trough. The next observation was that the yield group was daily exposed to social psychological stress. For instance, food intake was a typical stress event, suggesting that limited food may be one of the important reasons for natural depression behaviors in cynomolgus monkeys, which also mimics the high risk of social psychological stress produced by human depression. Finally, an interesting observation was that weak individuals adopted friendly behaviors such as grooming to avoid being attacked and promote social harmony. Further, the yield group tended to seek self-protection by warming together and forming “weak-weak combinations” to reduce attacks by other individuals [ 19 ]. Overall, group resources are limited, and social hierarchy is more strict. Some individuals are in such a chronic unpredictability of the stimulus state under the effect of such daily random conflict events [ 20 ]. This stimulus causes an increase in stress with time and ultimately leads to depression behaviors. Additionally, the high-level (attack) group was reported to be more active in conflict and motor behaviors [ 21 – 22 ]. This activity means that cynomolgus monkeys maintain their dominant state through attack behaviors. The low-level yield group showed stronger depression and anxious behaviors, which is similar to the finding that cynomolgus monkeys in the disadvantaged state are susceptible to developing depression behaviors [ 23 ]. The living environment and individuals are important factors affecting the expression of behavioral items in different groups. Still, the causes of depression behaviors are more consistent. Attacked individuals are unable to shift the pressure due to the chronic and unpredictable stimuli under random conflicts, which accumulate over time and eventually lead to depression behavior development. Some weak individuals prefer friendly behaviors as a communication channel, while others prefer warming behaviors to maintain their harmony. These behaviors also illustrate the diversity and complexity of the living environment of cynomolgus monkey communities. Additionally, the natural depression behaviors of cynomolgus monkeys are not accidental but are the result of intra-community conflicts. We designed a food exploration test and interactive behavioral evaluation and found that low-level cynomolgus monkeys spent more time exploring food than high-level ones. This difference indicated that the exploration capability of low-level individuals decreased significantly, which was similar to the embedded behavioral evaluation reported in a rodent animal model of depression [ 24 ]. Simultaneously, it was also found that the time for low-level individuals in the effective contact area within 5 min was significantly shortened in the interactive behavioral evaluation [ 25 ]. This observation suggests that the low-level group was more isolated and lacked social contact, which was similar to the condition of depression patients. Although the natural depression monkey model exhibited core behaviors, such as eye-opening, crouching, and head bowing, which showed its authenticity, the model screening did not include corresponding evaluations and verification. Hence, we designed the exploration and interaction experiment and found that cynomolgus monkey individuals with depression behaviors exhibited drastically degraded capability in exploration and social interaction capabilities. Consequently, it is reasonable to believe that the monkey models of natural depression screened in future research can provide corresponding behavioral evaluations to significantly increase the reliability and accuracy of these models, which is also the purpose of this study. Notwithstanding these findings, this study had several limitations. Firstly, the sample size was small, and individuals had statistically significant behaviors. Secondly, this report was a behavioral study, and non-human primate models of natural depression are not yet fully developed. Hence, future research needs to expand the research population, increase the number of validation experiments, and include non-human primate models of depression to improve observation and analysis methods. Conclusions In summary, this study investigated the causes of spontaneous depression in cynomolgus monkey communities, which have highly similar human social attributes to provide a reliable theoretical basis for the spontaneous depression model [ 26 ]. The subjects in this model lived in free developmental groups without any human intervention, truly reflecting the process of depression behaviors compared to other models. Simultaneously, the validation of depression behaviors in cynomolgus monkeys with depression behaviors was also conducted. The natural depression model in cynomolgus monkeys may be attributed to social defeat responses triggered by aggressive-yielding behaviors between groups. Abbreviations WHO World Health Organization DS The David Scoring AAALAC Association For Assessment and Accreditation of Laboratory Animal Care Declarations Ethics approval and consent to participate This study was approved by the Animal Ethics Committee of Chongqing Hospital of Traditional Chinese Medicine and followed the rules for the care and use of animals. Consent for publication Not Applicable Data availability The datasets used and analyzed during the current study are available from the corresponding author upon reasonable request. Competing interests The authors declare no competing interests. Funding This work was supported by the:Chongqing Ba Yu Qihuang Scholars Support Project (Yuzhongyi (2023) No. 23) . Authors’ contributions W.L. jointly conceived and designed the study.Y.H. collected the data; analyzed the data. S.G.L. drafted the original manuscript. J.T. supervised the data analysis and edited the manuscript for intellectual content and style. All authors read and approved the final version of the manuscript. Acknowledgments We express our gratitude to Zhongke Experimental Animal Co. Ltd., China for providing the facilities necessary to conduct our field observations. References Pitsillou E, Bresnehan SM, Kagarakis EA, Wijoyo SJ, Karagiannis TC. The cellular and molecular basis of major depressive disorder: towards a unified model for understanding clinical depression. Mol Biol Rep. 2020;47(1). Willner P. Validity, reliability and utility of the chronic mild stress model of depression: a 10-year review and evaluation. Psychopharmacology. 1997;134(4):319–29. Katz RJ, Roth KA, Carroll BJ. Acute and chronic stress effects on open field activity in the rat: implications for a model of depression. Neurosci Biobehavioral Reviews. 1981;5(2):247–51. Rosenblum LA, Paully GS. The effects of varying environmental demands on maternal and infant behavior. Child Dev. 1984;55(1):305–14. Shively CA, Register TC, Friedman DP, Morgan TM, Thompson J, Lanier T. Social stress-associated depression in adult female cynomolgus monkeys (Macaca fascicularis). Biol Psychol. 2005;69(1):67–84. Willard SL, Shively CA. Modeling depression in adult female cynomolgus monkeys (macaca fascicularis). Am J Primatol. 2012;74(6):528–42. Qin D, Rizak J, Chu X, Li Z, Yang S, Lv L, et al. A spontaneous depressive pattern in adult female rhesus macaques. Sci Rep. 2015;5:11267. Zhang ZY, Mao Y, Feng XL, et al. Early adversity contributes to chronic stress induced depression-like behavior in adolescent male rhesus monkeys. Behav Brain Res. 2016;306:154–9. Altmann J. Observational study of behavior: sampling methods. Behaviour. 1974;49(3):227–67. Noldus LPJJ. The observer: a software system for collection and analysis of observational data. Behav Res Methods Instruments Computers. 1991;23(3):415–29. Li X, Xu F, Xie L, et al. Depression-like behavioral phenotypes by social and social plus visual isolation in the adult female Macaca fascicularis. PLoS ONE. 2013;8(9):e73293. Xu F, Li X, Xie L, et al. Construction and validation of a systematic ethogram of Macaca fascicularis in a free enclosure. PLoS ONE. 2012;7(5):e37486. Gammell MP, Vries HD, Jennings DJ, Carlin CM, Hayden T. J Anim Behav. 2003;66:601–5. Chase ID. Models of hierarchy formation in animal societies. Syst Res Behav Sci. 1974;19(6):374–82. Simpson MJ, Simpson AE. Birth sex ratios and social rank in rhesus monkey mothers. Nature. 1982;300(5891):440–1. Worlein JM. Nonhuman primate models of depression: effects of early experience and stress. ILAR J. 2014;(2): 259. Shively CA, Laber-Laird K, Anton RF. Behavior and physiology of social stress and depression in female cynomolgus monkeys. Biol Psychiatry. 1997;41(8):871–82. Boldizsár Czéh, Fuchs E, Wiborg O. Mária Simon. Prog Neuro-Psychopharmacol Biol Psychiatry. 2016;64:293–310. Takahashi H. Changes of dominance rank, age, and tenure of wild japanese macaque males in the kinkazan a troop during seven years. J Primatol. 2002;43(2):13. Qin D, Chu X, Feng X, Li Z, Yang S, Lv L, et al. The first observation of seasonal affective disorder symptoms in Rhesus macaque. Behav Brain Res. 2015;292:463–9. Scott J, Lockard JS. Captive female gorilla agonistic relationships with clumped defendable food resources. Primates. 2006;47(3):199–209. Michels AM. Sex differences in food acquisition and aggression in captive common marmosets (callithrix jacchus). Primates. 1998;39(4):549. Shively CA, Kaplan JR, Adams MR. Effects of ovariectomy, social instability and social status on female Macaca fascicularis social behavior. Physiol Behav. 1986;36(6):1147–53. Wu R, Zhang H, Xue W, Zou Z, Chen G. Transgenerational impairment of hippocampal Akt-mTOR signaling and behavioral deficits in the offspring of mice that experience postpartum depression-like illness. Progress Neuropsychopharmacol Biol Psychiatry. 2017;73:11–8. Qiu-Qin H, Liu Y, Hui-Jie H, et al. Differential gr expression and translocation in the hippocampus mediates susceptibility vs. resilience to chronic social defeat stress. Front NeuroSci. 2017;11:287. Mansouri FA, Egner T, Buckley MJ. Monitoring demands for executive control:Shared functions between human and nonhuman primates. Trends Neurosci. 2017;40(1):15–27. Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 16 Oct, 2025 Read the published version in BMC Zoology → Version 1 posted Editorial decision: Revision requested 11 Jul, 2025 Reviews received at journal 10 Jul, 2025 Reviews received at journal 06 Jul, 2025 Reviewers agreed at journal 04 Jul, 2025 Reviewers agreed at journal 03 Jul, 2025 Reviewers agreed at journal 18 Jun, 2025 Reviewers invited by journal 18 Jun, 2025 Editor assigned by journal 18 Jun, 2025 Editor invited by journal 27 May, 2025 Submission checks completed at journal 26 May, 2025 First submitted to journal 26 May, 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. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6716220","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":473273454,"identity":"546c5cf1-b154-455c-908f-1652c3c69702","order_by":0,"name":"Wei Li","email":"","orcid":"","institution":"Chongqing Hospital of Traditional Chinese Medicine","correspondingAuthor":false,"prefix":"","firstName":"Wei","middleName":"","lastName":"Li","suffix":""},{"id":473273455,"identity":"c80e63e9-ace9-4e0d-a765-6c84bdbc4af7","order_by":1,"name":"Li Huang","email":"","orcid":"","institution":"Chongqing Hospital of Traditional Chinese Medicine","correspondingAuthor":false,"prefix":"","firstName":"Li","middleName":"","lastName":"Huang","suffix":""},{"id":473273456,"identity":"8841b881-4364-439e-8d51-d6896e03a7f1","order_by":2,"name":"Shi-Gang Liu","email":"","orcid":"","institution":"Non-human primates laboratory of Suzhou Zhongke Animal Experiment Co., Ltd","correspondingAuthor":false,"prefix":"","firstName":"Shi-Gang","middleName":"","lastName":"Liu","suffix":""},{"id":473273457,"identity":"d7d52e31-45b3-4368-b540-6176393cebba","order_by":3,"name":"Jun Tang","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA30lEQVRIiWNgGAWjYLACCQYGOQTvAAHVPFAtxkCKsYF4LUCQ2EC0Fnv2s4dfWLbdSe9vT37+4GcOgxzfjQTGzwX4bOHJS7OQbHuWO+PMM8PG3m0MxpI3EpilZ+B1WI6ZgWTb4dwNEjmMzYzbGBI33EhgY+bBp4X/DVhLugFUSz1hLRI5xg+AWhJgWhIMCGq58caMQeLcYUOQX2b2bpMwnHnmYbM0Pi3s/TnGnyXKDsvztyc/+PBzm4083/Hkg5/xaQECNmkJMJ0AIkBMaPzgAcwfPyC0jIJRMApGwSjABAC5wEsOpw9lnQAAAABJRU5ErkJggg==","orcid":"","institution":"Chongqing Hospital of Traditional Chinese Medicine","correspondingAuthor":true,"prefix":"","firstName":"Jun","middleName":"","lastName":"Tang","suffix":""}],"badges":[],"createdAt":"2025-05-21 11:53:22","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6716220/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6716220/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1186/s40850-025-00242-2","type":"published","date":"2025-10-16T15:57:34+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":85055393,"identity":"739ef832-23dd-48f0-baa4-dad6c25b6483","added_by":"auto","created_at":"2025-06-20 12:42:30","extension":"jpeg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":153330,"visible":true,"origin":"","legend":"\u003cp\u003eThe Macaca fascicularis of food exploration experiment\u003c/p\u003e\n\u003cp\u003eA:Feeding trough. B:Small rock fragment.C:food.\u003c/p\u003e","description":"","filename":"floatimage1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-6716220/v1/1d0ef1d0d8e407a9271fb66e.jpeg"},{"id":85054478,"identity":"aa3bd9ea-92cf-482d-ad28-96d14132a56f","added_by":"auto","created_at":"2025-06-20 12:34:30","extension":"jpeg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":431575,"visible":true,"origin":"","legend":"\u003cp\u003eThe Macaca fascicularis of social interaction test\u003c/p\u003e\n\u003cp\u003eA:Transparent partition（The region extending 0.5 meters to the left and right of the transparent partition is defined as the effective contact zone).\u003c/p\u003e","description":"","filename":"floatimage2.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-6716220/v1/34ed131cd7e295deeadbc02e.jpeg"},{"id":93955984,"identity":"1c80dee0-2bbf-48d9-b733-0535035d4b81","added_by":"auto","created_at":"2025-10-20 16:08:51","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1834422,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6716220/v1/af659bf8-7160-48f1-a570-635aca557c59.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Investigation and validation of a natural depression model in cynomolgus monkeys （Macaca fascicularis）","fulltext":[{"header":"Background","content":"\u003cp\u003eAccording to 2019 statistics released by the World Health Organization (WHO), 320 million depression cases were reported globally by 2019, of which 95\u0026nbsp;million depression cases were reported in China. Furthermore, research has shown that depression ranks second among the largest disease burdens in China [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. There are no clear experimental indicators because of the inherent limitations of human studies and the unclear pathogenesis of depression. Hence, the use of animal models remains crucial in identifying the neural mechanisms of depression and discovering new drugs and treatments. Currently, rodent animal models, such as the chronic unpredictable mild stimulus model, behavioral despair model, and the social frustration stress model, are used in basic depression research [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. However, these animal models are established in non-social environments. However, depression is a disease closely related to social pressure and social relationships. Further, rodent animal models have several limitations, including the lack of complex characteristics seen in human society. Non-human primate models of depression can compensate for this challenge of rodent systems. Currently, non-human primate models of depression include the social isolation model, mother-infant separation model, and natural depression model [\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e]. Among these models, the natural depression model is the most ethical and socially relevant one. The approach identifies animals with eye-opening, crouching, and head bowing as core behaviors in a caged community of cynomolgus monkeys (Macaca fascicularis) [\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]. For instance, Shively et al. confirmed a few monkeys with natural depression, although the causes of natural depression in the animals were not specified [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e]. Further, even when this natural depression model was screened, it was not validated. Given these findings, a community of cynomolgus monkeys was established by mimicking wild conditions. The social behaviors of cynomolgus monkeys were recorded, and the causes of their depression-like behaviors were investigated. Further, the interactive behavior and food exploration behavior were evaluated to validate the results. This study aims to examine the causes of natural depression behaviors and validate them with corresponding evaluations. The results can provide reliable theoretical support for future studies of cynomolgus monkey models of natural depression.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eAnimal details and selection\u003c/h2\u003e \u003cp\u003e This study was conducted at the non-human primates laboratory of Suzhou Zhongke Animal Experiment Co., Ltd, certified by the Association For Assessment and Accreditation of Laboratory Animal Care (AAALAC). The cynomolgus monkeys were provided by the Zhongke Animal Experimental Base. The license number for the domestication and breeding of nationally protected wild animals was SuFa Domestication 2007-001. Initially, 22 healthy cynomolgus monkeys from different cages were randomly selected from the Zhongke Animal Experimental Base. After excluding pregnancy and other interfering factors, 19 cynomolgus monkeys were selected to form a community. The average age was 8.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1 years, and the male-to-female ratio was 1:8.5. The housing met the cage standards for caged cynomolgus monkeys in China. The basic data of the cynomolgus monkeys in this study is shown in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe basic data of the cynomolgus monkeys(\u003cem\u003eMacaca fascicularis\u003c/em\u003e)\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"8\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNumber\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eExperiment No.\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eGender\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAge (years)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNumber\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eExperiment No.\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eGender\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eAge (years)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e12152\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e06121\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e11192\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e05117\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e08264\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e01102\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e03150\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e11240\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e12052\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e03312\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e04126\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e07258\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e05056\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e05128\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e01140\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e07162\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e03128\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e12118\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e08198\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eMethods\u003c/h3\u003e\n\u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eEstablishment of the cynomolgus monkey community\u003c/h2\u003e \u003cp\u003eCynomolgus monkeys may be exposed to fierce conflict behaviors due to competition for food resources, territory, and mating rights [\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]. In this study, a community of \u0026ldquo;multiple females and multiple males\u0026rdquo; was created by simulating the wild living habitats of cynomolgus monkeys.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eObserving the daily behaviors and conflict behaviors\u003c/h3\u003e\n\u003cp\u003eThe characteristics of cynomolgus monkey individuals were identified in the pre-observation phase conducted for 2 weeks to provide an accurate basis for the subsequent recording of various behaviors. After the pre-observation period, each community of cynomolgus monkeys was assigned four experimental recorders; two were observed for attack-yield behaviors, and the other two were photographed and observed for daily behaviors. Different behaviors were observed in ten time slots, and a consistency test was performed among the behaviors, with k\u0026thinsp;\u0026gt;\u0026thinsp;80% as valid observations. The observation researchers recorded the effective number of attack-yield behaviors according to the behavioral recording items. The researchers who observed the daily behavior followed and photographed cynomolgus monkeys using the focused filming method [\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e] .The observation time was 8:30\u0026thinsp;\u0026minus;\u0026thinsp;11:30 and 14:00\u0026ndash;16:00; each time slot was 30 min, with ten time slots per day, and the cumulative effective time was 1050 h. The Noldus analysis system was used to analyze animal behavior [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e].\u003c/p\u003e\n\u003ch3\u003eRecord of conflict behaviors\u003c/h3\u003e\n\u003cp\u003eThe conflict behaviors are defined in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e based on the behavioral spectrum of cynomolgus monkeys [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e].\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eDefinition of cynomolgus monkeys(\u003cem\u003eMacaca fascicularis\u003c/em\u003e) Conflict Behavior\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"3\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBehavior category\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBehavioral definition\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAttack behaviour\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBite\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAggressive behaviors such as biting each other's head, back, limbs, and other body parts with teeth represent the most severe form of aggression, which can result in fatal consequences for the attacked individual\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGrab, hit, push\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMinimal physical contact, infrequently resulting in bodily harm\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eThreats\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEngaging in verbal altercations and forcefully shaking doors and windows with upper limbs\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYielding behavior\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eRun away\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eQuickly flee from the attacker after being attacked and scream.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePassive submissio\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFollowing an attack, the individual does not flee from the aggressor, instead emitting a scream and displaying friendly behaviors towards the attacker, such as grooming\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAvoidance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eThe two parties do not engage in physical confrontation and deliberately distance themselves from their original positions\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eDefinition and records of daily behaviors\u003c/h2\u003e \u003cp\u003eThe animal behavior analysis system was used to record and analyze ten types of behaviors, including anxiety, conflicts, and food intake [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]. Friendly behaviors refer to individuals grooming other individuals after an active attack or being attacked. Anxious behaviors are noted in individuals who rapidly scratch for more than 3 s. Rest behaviors refer to standing, sitting, or lying down for more than 5 s. Conflict behaviors include biting, scratching, or threatening another individual. Food intake behaviors refer to individuals grabbing food from the trough. Warming behaviors refer to individuals snuggling with each other to maintain body temperature. Mating behaviors refer to sexual behaviors between individuals of opposite sexes. Motor behaviors include moving quickly or slowly, with a distance greater than half an arm\u0026rsquo;s length. Depression behaviors refer to the presence of core depression-like behaviors, including eye-opening, crouching, and head- bowing. Finally, other behaviors refer to behaviors other than those in the above nine categories.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eFood exploration test\u003c/h3\u003e\n\u003cp\u003eIndividuals were divided into high- and low-level groups based on their David Scoring (DS) scores. Food (apples) was placed in the trough of a separate cage (2m \u0026times;1m \u0026times;1m) and buried with stones. A high-definition camera was placed 1 m in front of the cage to record the behavior of the cynomolgus monkeys with the apples without any interference. After turning on the camera, the experimenters left the cage, and recorded the time from placing the food to the time when the subject touched the food for each monkey. The subject animal was familiarized with the cage environment before the experiment, as shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eInteractive behavioral evaluation\u003c/h3\u003e\n\u003cp\u003eSingle cages (2m\u0026times;1m\u0026times;1m) were separated from each other by a transparent plastic partition with holes of 1 cm in diameter to avoid contact between subjects but not to isolate them from sight and smell. Further, high- and low-level groups from each community were tested separately. The interaction subject was a healthy male cynomolgus monkey from another community, and the effective contact area was defined as the area 0.5 m to the left and right of the transparent partition. Each experiment lasted 5 min, and the time spent in the effective contact area by each cynomolgus monkey individual within 5 min from placing the male cynomolgus monkeys was recorded, as shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eData on the daily behavior indicators were expressed as means\u0026thinsp;\u0026plusmn;\u0026thinsp;\u003cem\u003eSD\u003c/em\u003e, and the Shapiro-Wilk test was used to test the normal distribution of the eleven behavioral aspects. As the data were not normally distributed, the nonparametric Wilcoxon rank sum test was used for the statistical analysis, and the significance level was set at α\u0026thinsp;=\u0026thinsp;0.05. The attack-yield indicators were expressed as times, and the DS scores of each subject were calculated based on the attack-yield relationship matrix using the David\u0026rsquo;s score method. The data of food exploration tests were expressed as means\u0026thinsp;\u0026plusmn;\u0026thinsp;\u003cem\u003eSD\u003c/em\u003e and an independent samples t-test was conducted, with the significance level atα\u0026thinsp;=\u0026thinsp;0.05.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eAttack-yield (conflict) behavior correlation matrix and David Scoring scores\u003c/h2\u003e \u003cp\u003eThe established communities had significantly intense conflict behaviors since their formation, and 1122 valid conflict behaviors were recorded. David Scoring (DS) method was used to count the DS scores of each subject, and the formula was DS\u0026thinsp;=\u0026thinsp;W1\u0026thinsp;+\u0026thinsp;W2 - L1 - L2. Here, W1 was the sum of the dominant percentages of monkeys who exhibited conflict behaviors in the group; W2 was the sum of the dominant weighted percentage in individual conflict behaviors; L1 was the sum of the disadvantage percentages of animals who exhibited conflict behaviors in the group; and L2 was the sum of the weighted disadvantage percentages in individual conflict behaviors [\u003cspan additionalcitationids=\"CR14\" citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e]. The DS scores are shown in Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e\u003cb\u003eMatrix of aggressive behavior relationships and individual DS scores for the group\u003c/b\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"24\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c14\" colnum=\"14\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c15\" colnum=\"15\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c16\" colnum=\"16\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c17\" colnum=\"17\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c18\" colnum=\"18\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c19\" colnum=\"19\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c20\" colnum=\"20\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c21\" colnum=\"21\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c22\" colnum=\"22\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c23\" colnum=\"23\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c24\" colnum=\"24\"\u003e\u003c/div\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c12\" namest=\"c10\"\u003e \u003cp\u003eAttacker\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c15\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eDS scores\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003eGenuflector\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eM\u003csub\u003e1\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eF\u003csub\u003e1\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eF\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eF\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eF\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e \u003cp\u003eF\u003csub\u003e5\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eF\u003csub\u003e6\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003eF\u003csub\u003e7\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003eF\u003csub\u003e8\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003eF\u003csub\u003e9\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003eF\u003csub\u003e10\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003eF\u003csub\u003e11\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003eF\u003csub\u003e12\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003eF\u003csub\u003e13\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003eF\u003csub\u003e14\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003eF\u003csub\u003e15\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003eF\u003csub\u003e16\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003eF\u003csub\u003e17\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" morerows=\"18\" nameend=\"c2\" namest=\"c1\" rowspan=\"19\"\u003e \u003cp\u003eM\u003csub\u003e1\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eM\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e1\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e5\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e6\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e7\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e8\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e9\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e10\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e11\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e12\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e13\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e14\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e15\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e16\u003c/sub\u003e\u003c/p\u003e \u003cp\u003eF\u003csub\u003e17\u003c/sub\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e171\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e152\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e131\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e116\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e95\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e73\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e57\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e-18\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e-38\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e-57\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e-88\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e-94\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e-119\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e-120\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e-150\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c13\" namest=\"c12\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c15\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c21\"\u003e \u003cp\u003e22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c22\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c23\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c24\"\u003e \u003cp\u003e-171\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eDaily behaviors of attack and yield groups\u003c/h2\u003e \u003cp\u003eIndividuals with positive and negative DS scores were divided into the high-level group (the attack group) and the low-level group (the yield group), respectively, and their daily behaviors were observed. The animals showed statistically significant differences in friendly, anxious, conflicting, mating, motor, and depression behaviors (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05), as shown in Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e\u003cb\u003eDaily Observations of the Aggressor Group and the Yielding Group\u003c/b\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBehavioral items\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAggressor Group\u003c/p\u003e \u003cp\u003e(n\u0026thinsp;=\u0026thinsp;10)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eYielding Group\u003c/p\u003e \u003cp\u003e(n\u0026thinsp;=\u0026thinsp;9)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003et\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eP\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDepression behaviors\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.01\u0026thinsp;\u0026plusmn;\u0026thinsp;1.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e30.44\u0026thinsp;\u0026plusmn;\u0026thinsp;23.90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-3.691\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.006*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFriendly behaviors\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e49.30\u0026thinsp;\u0026plusmn;\u0026thinsp;24.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e140.77\u0026thinsp;\u0026plusmn;\u0026thinsp;48.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-5.085\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.024*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMating behaviors\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4.88\u0026thinsp;\u0026plusmn;\u0026thinsp;1.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.88\u0026thinsp;\u0026plusmn;\u0026thinsp;1.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.731\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.040*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eConflict behaviors\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e76.30\u0026thinsp;\u0026plusmn;\u0026thinsp;22.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e12.55\u0026thinsp;\u0026plusmn;\u0026thinsp;7.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.478\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.018*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRest behaviors\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e800.60\u0026thinsp;\u0026plusmn;\u0026thinsp;81.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e709.00\u0026thinsp;\u0026plusmn;\u0026thinsp;50.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.663\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.518\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWarming behaviors\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e23.50\u0026thinsp;\u0026plusmn;\u0026thinsp;6.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e22.44\u0026thinsp;\u0026plusmn;\u0026thinsp;5.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.503\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.551\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAnxious behaviors\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e11.20\u0026thinsp;\u0026plusmn;\u0026thinsp;8.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e73.11\u0026thinsp;\u0026plusmn;\u0026thinsp;73.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-4.053\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.005*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMotor behaviors\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e141.1\u0026thinsp;\u0026plusmn;\u0026thinsp;25.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e31.77\u0026thinsp;\u0026plusmn;\u0026thinsp;24.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e9.639\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.007*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFood intake behaviors\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e48.60\u0026thinsp;\u0026plusmn;\u0026thinsp;14.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7.88\u0026thinsp;\u0026plusmn;\u0026thinsp;6.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.284\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.780\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOther behaviors\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e47.00\u0026thinsp;\u0026plusmn;\u0026thinsp;12.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e50.11\u0026thinsp;\u0026plusmn;\u0026thinsp;22.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.382\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.707\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"5\"\u003eData are means\u0026thinsp;\u0026plusmn;\u0026thinsp;\u003cem\u003eSD\u003c/em\u003e. \u003cem\u003e*\u003c/em\u003e: \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eFood exploration tests\u003c/h2\u003e \u003cp\u003eThe time taken by the attack and yield groups to reach the food after placing apples was recorded. The results were statistically significant (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05), suggesting that low-level (yield) cynomolgus monkeys spent more time exploring the food than high-level (attack) ones (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e\u003cb\u003eComparison of the time taken by individuals of varying hierarchical positions within a group to explore food\u003c/b\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAggressor Group\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eYielding Group\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003et\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eP\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003en\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTime taken to search for and find food\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.10\u0026thinsp;\u0026plusmn;\u0026thinsp;2.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e15.03\u0026thinsp;\u0026plusmn;\u0026thinsp;4.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.422\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.015*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"5\"\u003eData are means\u0026thinsp;\u0026plusmn;\u0026thinsp;\u003cem\u003eSD\u003c/em\u003e. \u003cem\u003e*\u003c/em\u003e: \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eThe social interaction test in cynomolgus monkeys\u003c/h2\u003e \u003cp\u003eThe time spent in the effective contact area within 5 minutes was recorded in high-level and low-level individuals in the group. The low-level (yield) individuals spent a significantly smaller time in the effective contact area than that of high-level (attack) monkeys, with statistically significant differences (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab6\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 6\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e\u003cb\u003eComparison of the time that individuals of various ranks within a group spend in the effective contact area\u003c/b\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAggressor Group\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eYielding Group\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003et\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eP\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003en\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTime spent within the effective contact area\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4.3\u0026thinsp;\u0026plusmn;\u0026thinsp;1.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.01\u0026thinsp;\u0026plusmn;\u0026thinsp;0.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.242\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.007\u003cem\u003e*\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"5\"\u003eData are means\u0026thinsp;\u0026plusmn;\u0026thinsp;\u003cem\u003eSD\u003c/em\u003e. \u003cem\u003e*\u003c/em\u003e: \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eAnimal models with depression have always been vital for depression research. Further, the pathogenesis of depression can be better investigated with their high similarity with depression symptoms [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]. Currently, most animal models for depression use rodents, which lack certain social relevance. This lack can be compensated using non-human primate models of depression for this more accurate studies. For instance, Shively reported that some individuals spontaneously exhibited human-like depression due to social pressure in cynomolgus monkey communities [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. However, the depression behaviors of cynomolgus monkeys are mainly reflected as crouching, eye-opening, and decreased mobility, which are different from those of humans. The cynomolgus monkey communities have strict social hierarchies, and the yield group is under long-term social pressure due to the lack of access to mating rights and food [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]. It is therefore hypothesized that these individuals are readily exposed to depression-like behaviors. In this study, we observed the attack-yield behaviors of cynomolgus monkeys after establishing a community. We recorded and analyzed the daily behaviors of individuals to explore the occurrence and the possible causes of depression-like behaviors in caged cynomolgus monkeys.\u003c/p\u003e \u003cp\u003eIn this study, 1122 behaviors were observed, and more intense attack-yield behaviors occurred from the beginning of group formation. Individuals were classified into the attack group and the yield group based on their DS scores, with significant differences in their daily behaviors. There were significant differences in the friendly, anxious, conflicting, mating, motor, and depression behaviors. The attack group was more likely to initiate conflicts, mating behavior, and motor behaviors with other individuals In contrast, the yield group was more likely to exhibit friendly, depressed, and anxious behaviors. The differences in group behaviors revealed the following characteristics: firstly, the attack group performed more prominently in attack behaviors than the yield group. The group attack-yield behavior correlation matrix showed that some community individuals could mitigate pressure by attacking and threatening other individuals when subjected to attacks and threats. In contrast, individuals with low DS scores had difficulty handling this pressure and were readily exposed to anxious and depressive behaviors. Further, the yield group was threatened by more individuals and rarely received friendly grooming behaviors from other individuals. The cynomolgus monkeys of the yield group developed more friendly behaviors to promote harmony with other individuals and avoid being attacked by others. Thirdly, as expected, the yield group developed typical solitary sitting, head bowing, crouching, and eating in more remote areas away from the trough. The next observation was that the yield group was daily exposed to social psychological stress. For instance, food intake was a typical stress event, suggesting that limited food may be one of the important reasons for natural depression behaviors in cynomolgus monkeys, which also mimics the high risk of social psychological stress produced by human depression. Finally, an interesting observation was that weak individuals adopted friendly behaviors such as grooming to avoid being attacked and promote social harmony. Further, the yield group tended to seek self-protection by warming together and forming \u0026ldquo;weak-weak combinations\u0026rdquo; to reduce attacks by other individuals [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]. Overall, group resources are limited, and social hierarchy is more strict. Some individuals are in such a chronic unpredictability of the stimulus state under the effect of such daily random conflict events [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]. This stimulus causes an increase in stress with time and ultimately leads to depression behaviors.\u003c/p\u003e \u003cp\u003eAdditionally, the high-level (attack) group was reported to be more active in conflict and motor behaviors [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]. This activity means that cynomolgus monkeys maintain their dominant state through attack behaviors. The low-level yield group showed stronger depression and anxious behaviors, which is similar to the finding that cynomolgus monkeys in the disadvantaged state are susceptible to developing depression behaviors [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]. The living environment and individuals are important factors affecting the expression of behavioral items in different groups. Still, the causes of depression behaviors are more consistent. Attacked individuals are unable to shift the pressure due to the chronic and unpredictable stimuli under random conflicts, which accumulate over time and eventually lead to depression behavior development. Some weak individuals prefer friendly behaviors as a communication channel, while others prefer warming behaviors to maintain their harmony. These behaviors also illustrate the diversity and complexity of the living environment of cynomolgus monkey communities. Additionally, the natural depression behaviors of cynomolgus monkeys are not accidental but are the result of intra-community conflicts.\u003c/p\u003e \u003cp\u003eWe designed a food exploration test and interactive behavioral evaluation and found that low-level cynomolgus monkeys spent more time exploring food than high-level ones. This difference indicated that the exploration capability of low-level individuals decreased significantly, which was similar to the embedded behavioral evaluation reported in a rodent animal model of depression [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. Simultaneously, it was also found that the time for low-level individuals in the effective contact area within 5 min was significantly shortened in the interactive behavioral evaluation [\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e]. This observation suggests that the low-level group was more isolated and lacked social contact, which was similar to the condition of depression patients. Although the natural depression monkey model exhibited core behaviors, such as eye-opening, crouching, and head bowing, which showed its authenticity, the model screening did not include corresponding evaluations and verification. Hence, we designed the exploration and interaction experiment and found that cynomolgus monkey individuals with depression behaviors exhibited drastically degraded capability in exploration and social interaction capabilities. Consequently, it is reasonable to believe that the monkey models of natural depression screened in future research can provide corresponding behavioral evaluations to significantly increase the reliability and accuracy of these models, which is also the purpose of this study.\u003c/p\u003e \u003cp\u003eNotwithstanding these findings, this study had several limitations. Firstly, the sample size was small, and individuals had statistically significant behaviors. Secondly, this report was a behavioral study, and non-human primate models of natural depression are not yet fully developed. Hence, future research needs to expand the research population, increase the number of validation experiments, and include non-human primate models of depression to improve observation and analysis methods.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eIn summary, this study investigated the causes of spontaneous depression in cynomolgus monkey communities, which have highly similar human social attributes to provide a reliable theoretical basis for the spontaneous depression model [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]. The subjects in this model lived in free developmental groups without any human intervention, truly reflecting the process of depression behaviors compared to other models. Simultaneously, the validation of depression behaviors in cynomolgus monkeys with depression behaviors was also conducted. The natural depression model in cynomolgus monkeys may be attributed to social defeat responses triggered by aggressive-yielding behaviors between groups.\u003c/p\u003e"},{"header":"Abbreviations","content":"\u003cp\u003eWHO \u0026nbsp; \u0026nbsp; \u0026nbsp;World Health Organization\u003c/p\u003e\n\u003cp\u003eDS \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;The David Scoring\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAAALAC \u0026nbsp;Association For Assessment and Accreditation of Laboratory Animal Care\u0026nbsp;\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis study was approved by the Animal Ethics Committee of Chongqing Hospital of Traditional Chinese Medicine and followed the rules for the care and use of animals.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eNot Applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets used and analyzed during the current study are available from the corresponding author upon reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by the:Chongqing Ba Yu Qihuang Scholars Support Project (Yuzhongyi (2023) No. 23) .\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors\u0026rsquo; \u0026nbsp;contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eW.L. jointly conceived and designed the study.Y.H. collected the data; analyzed the data. S.G.L. drafted the original manuscript. J.T. supervised the data analysis and edited the manuscript for intellectual content and style. All authors read and approved the final version of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe express our gratitude to Zhongke Experimental Animal Co. Ltd., China for providing the facilities necessary to conduct our field observations.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003ePitsillou E, Bresnehan SM, Kagarakis EA, Wijoyo SJ, Karagiannis TC. The cellular and molecular basis of major depressive disorder: towards a unified model for understanding clinical depression. Mol Biol Rep. 2020;47(1).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWillner P. Validity, reliability and utility of the chronic mild stress model of depression: a 10-year review and evaluation. Psychopharmacology. 1997;134(4):319\u0026ndash;29.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKatz RJ, Roth KA, Carroll BJ. Acute and chronic stress effects on open field activity in the rat: implications for a model of depression. Neurosci Biobehavioral Reviews. 1981;5(2):247\u0026ndash;51.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRosenblum LA, Paully GS. The effects of varying environmental demands on maternal and infant behavior. Child Dev. 1984;55(1):305\u0026ndash;14.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eShively CA, Register TC, Friedman DP, Morgan TM, Thompson J, Lanier T. Social stress-associated depression in adult female cynomolgus monkeys (Macaca fascicularis). Biol Psychol. 2005;69(1):67\u0026ndash;84.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWillard SL, Shively CA. Modeling depression in adult female cynomolgus monkeys (macaca fascicularis). Am J Primatol. 2012;74(6):528\u0026ndash;42.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eQin D, Rizak J, Chu X, Li Z, Yang S, Lv L, et al. A spontaneous depressive pattern in adult female rhesus macaques. Sci Rep. 2015;5:11267.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang ZY, Mao Y, Feng XL, et al. Early adversity contributes to chronic stress induced depression-like behavior in adolescent male rhesus monkeys. Behav Brain Res. 2016;306:154\u0026ndash;9.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAltmann J. Observational study of behavior: sampling methods. Behaviour. 1974;49(3):227\u0026ndash;67.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eNoldus LPJJ. The observer: a software system for collection and analysis of observational data. Behav Res Methods Instruments Computers. 1991;23(3):415\u0026ndash;29.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLi X, Xu F, Xie L, et al. Depression-like behavioral phenotypes by social and social plus visual isolation in the adult female Macaca fascicularis. PLoS ONE. 2013;8(9):e73293.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eXu F, Li X, Xie L, et al. Construction and validation of a systematic ethogram of Macaca fascicularis in a free enclosure. PLoS ONE. 2012;7(5):e37486.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGammell MP, Vries HD, Jennings DJ, Carlin CM, Hayden T. J Anim Behav. 2003;66:601\u0026ndash;5.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChase ID. Models of hierarchy formation in animal societies. Syst Res Behav Sci. 1974;19(6):374\u0026ndash;82.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSimpson MJ, Simpson AE. Birth sex ratios and social rank in rhesus monkey mothers. Nature. 1982;300(5891):440\u0026ndash;1.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWorlein JM. Nonhuman primate models of depression: effects of early experience and stress. ILAR J. 2014;(2): 259.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eShively CA, Laber-Laird K, Anton RF. Behavior and physiology of social stress and depression in female cynomolgus monkeys. Biol Psychiatry. 1997;41(8):871\u0026ndash;82.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBoldizs\u0026aacute;r Cz\u0026eacute;h, Fuchs E, Wiborg O. M\u0026aacute;ria Simon. Prog Neuro-Psychopharmacol Biol Psychiatry. 2016;64:293\u0026ndash;310.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTakahashi H. Changes of dominance rank, age, and tenure of wild japanese macaque males in the kinkazan a troop during seven years. J Primatol. 2002;43(2):13.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eQin D, Chu X, Feng X, Li Z, Yang S, Lv L, et al. The first observation of seasonal affective disorder symptoms in Rhesus macaque. Behav Brain Res. 2015;292:463\u0026ndash;9.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eScott J, Lockard JS. Captive female gorilla agonistic relationships with clumped defendable food resources. Primates. 2006;47(3):199\u0026ndash;209.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMichels AM. Sex differences in food acquisition and aggression in captive common marmosets (callithrix jacchus). Primates. 1998;39(4):549.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eShively CA, Kaplan JR, Adams MR. Effects of ovariectomy, social instability and social status on female Macaca fascicularis social behavior. Physiol Behav. 1986;36(6):1147\u0026ndash;53.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWu R, Zhang H, Xue W, Zou Z, Chen G. Transgenerational impairment of hippocampal Akt-mTOR signaling and behavioral deficits in the offspring of mice that experience postpartum depression-like illness. Progress Neuropsychopharmacol Biol Psychiatry. 2017;73:11\u0026ndash;8.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eQiu-Qin H, Liu Y, Hui-Jie H, et al. Differential gr expression and translocation in the hippocampus mediates susceptibility vs. resilience to chronic social defeat stress. Front NeuroSci. 2017;11:287.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMansouri FA, Egner T, Buckley MJ. Monitoring demands for executive control:Shared functions between human and nonhuman primates. Trends Neurosci. 2017;40(1):15\u0026ndash;27.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"bmc-zoology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"bzoo","sideBox":"Learn more about [BMC Zoology](http://bmczool.biomedcentral.com/)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/bzoo/default.aspx","title":"BMC Zoology","twitterHandle":"@BMC_series","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"em","reportingPortfolio":"BMC Series","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"MDD, Macaca fascicularis, Chronic unpredictable mild stress, Social interaction test","lastPublishedDoi":"10.21203/rs.3.rs-6716220/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6716220/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eBackground\u003c/h2\u003e \u003cp\u003eThe global burden of depression is heavy, with unclear pathogenesis, relying on animal models for researching neural mechanisms and treatments. Existing rodent models lack social attributes, while non-human primate models (such as the natural depression model in cynomolgus monkeys) are closer to human social stress scenarios. Previously, the screening of natural depression models in cynomolgus monkeys was unvalidated and their causes were unclear. This study constructs a cynomolgus monkey community by simulating wild environments, records social behaviors, and explores the causes of their natural depression-like behaviors and validates them, providing theoretical support for this model.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e \u003cp\u003eThe conflict behaviors in the community were recorded to develop a conflict behaviors matrix, and the behaviors of all individuals were monitored by using the focus observation method. The analysis entailed recording and analyzing ten behaviors for ten observation time slots on individuals with depression. The David Scoring (DS) scores of the animals were determined based on the record of conflict behaviors in the community. The individuals were divided into high- and low-level groups, and the daily behaviors of the two groups were analyzed. The results showed statistically significant differences in depression behaviors, anxious behaviors, and motor behaviors among the monkeys. Further, individuals with depression behaviors exhibited drastically lower capability in exploration and social interaction. Additionally, social frustration stress responses induced by inter- and intra-community conflict behaviors may account for this behavior in the cynomolgus monkey model of natural depression.\u003c/p\u003e\u003ch2\u003eConclusions\u003c/h2\u003e \u003cp\u003eThe natural depression model of caged Macaca fascicularis is a social defeat model induced by yield-attack behavior.\u003c/p\u003e","manuscriptTitle":"Investigation and validation of a natural depression model in cynomolgus monkeys （Macaca fascicularis）","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-20 12:26:25","doi":"10.21203/rs.3.rs-6716220/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-07-11T12:59:28+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-10T07:08:02+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-06T23:59:39+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"307463882460282602115957620289184720759","date":"2025-07-04T12:51:04+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"67219972557875956056585765053914241294","date":"2025-07-04T00:30:51+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"186974633143162630019910604443586005126","date":"2025-06-18T14:37:00+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-18T14:30:00+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-06-18T08:45:34+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2025-05-27T10:48:37+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-05-26T13:59:09+00:00","index":"","fulltext":""},{"type":"submitted","content":"BMC Zoology","date":"2025-05-26T13:58:00+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"bmc-zoology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"bzoo","sideBox":"Learn more about [BMC Zoology](http://bmczool.biomedcentral.com/)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/bzoo/default.aspx","title":"BMC Zoology","twitterHandle":"@BMC_series","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"em","reportingPortfolio":"BMC Series","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"99189025-0337-4d3a-9a0f-653a51c4b912","owner":[],"postedDate":"June 20th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-10-20T16:01:25+00:00","versionOfRecord":{"articleIdentity":"rs-6716220","link":"https://doi.org/10.1186/s40850-025-00242-2","journal":{"identity":"bmc-zoology","isVorOnly":false,"title":"BMC Zoology"},"publishedOn":"2025-10-16 15:57:34","publishedOnDateReadable":"October 16th, 2025"},"versionCreatedAt":"2025-06-20 12:26:25","video":"","vorDoi":"10.1186/s40850-025-00242-2","vorDoiUrl":"https://doi.org/10.1186/s40850-025-00242-2","workflowStages":[]},"version":"v1","identity":"rs-6716220","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6716220","identity":"rs-6716220","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}