No evidence for a link between maternal aggression and inhibitory control across generations in domestic canaries

No evidence for a link between maternal aggression and inhibitory control across generations in domestic canaries | 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 No evidence for a link between maternal aggression and inhibitory control across generations in domestic canaries Clara Garcia-Co, Anneleen Dewulf, Frederick Verbruggen, Judith Morales, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9402003/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 5 You are reading this latest preprint version Abstract Maternal aggression (MA) towards offspring represents an early-life social stressor and has been linked to long-term behavioural differences. Yet, the cognitive mechanisms underlying its expression and potential intergenerational transmission remain unclear. We tested whether MA in domestic canaries ( Serinus canaria ) is associated with individual differences in inhibitory control (i.e., the ability to suppress impulsive or inappropriate actions) in adult females and whether early-life exposure to MA subsequently predicts variation in offspring inhibitory control. Inhibitory control was assessed using a detour task, in which individuals must suppress a direct approach to a visible reward in favour of an indirect solution. A cross-fostering design was used to disentangle post-hatching environmental effects from prenatal or genetic influences on offspring performance. We found no evidence that aggressive and non-aggressive mothers differed in inhibitory control. Likewise, offspring inhibitory control was not influenced by post-hatching exposure to MA nor by the MA phenotype of the biological mother, and we detected no mother–offspring resemblance in inhibitory control. However, strong ontogenetic effects were observed: juveniles showed longer detour latencies than adults but improved more steeply across trials, consistent with a post-hatching maturation of inhibitory control. Together, these results suggest that MA in this system is unlikely to reflect deficits in inhibitory control or to be transmitted across generations via this cognitive pathway. More broadly, our findings highlight the task- and context-specific nature of inhibitory control measures. inhibitory control response inhibition maternal aggression detour task transgenerational transmission ontogeny Figures Figure 1 Figure 2 Figure 3 Figure 4 INTRODUCTION Parental care is a fundamental component of the offspring's early-life environment and plays a crucial role in shaping cognitive and behavioural development. Even though both sexes may contribute to parental care in most species, often mothers have a greater impact, as they can influence offspring development to a larger extent both prenatally, e.g., through hormone deposition in eggs, and postnatally, through more frequent behavioural interactions during which they provide resources, protection, and social cues (Bagot & Meaney, 2010 ; Crews, 2010 ; Groothuis et al., 2005 ; Reddon, 2012 ). These maternal effects are increasingly recognised as key drivers of individual differences in behaviour and cognition, which can persist across life stages (Bagot & Meaney, 2010 ; McEwen & Morrison, 2013 ; Meaney, 2010 ; Veenema, 2009 ). While maternal care is typically beneficial, maternal behaviour can also include aggressive interactions directed towards the offspring. Maternal aggression (MA), such as pecking, chasing, or attacking offspring, has been documented across taxa and is often expressed under conditions of environmental stress, high competition, or reproductive conflict (Arnold & Taborsky, 2010 ; Champagne & Meaney, 2007 ; Garcia-Co et al., 2025 ; Leonard et al., 1988 ; Maestripieri, 1998 ; Shizuka & Lyon, 2013 ). When occurring during sensitive periods of offspring development, MA has the potential to shape offspring development too, although likely adversely. This raises important questions about the causation and consequences of MA (Anda et al., 2006 ; Arnold & Taborsky, 2010 ; Bannier et al., 2017 ; Gondré-Lewis et al., 2016 ; Lovallo, 2013 ). One potential causal mechanism associated with the expression of MA is impaired inhibitory control. Inhibitory control allows individuals to suppress impulsive or inappropriate actions; it is a core executive function that underpins goal-directed and adaptive behaviours (Diamond, 2013 ; Verbruggen & Logan, 2008 ). In particular, response inhibition refers to the ability to suppress a prepotent motor response (Aron, 2011 ; Verbruggen & Logan, 2008 ). Impairments in response inhibition have been linked to reactive forms of aggression, which are characterised by rapid, emotionally driven responses to perceived challenges (Berkowitz, 1993 ; Feilhauer et al., 2012 ; Hecht & Latzman, 2018 ; Zhang et al., 2017 ). Across taxa, reduced response inhibition and heightened impulsivity have been associated to altered functioning of prefrontal circuits involved in behavioural regulation (Madole et al., 2020 ; Puiu et al., 2018 ; Rudebeck et al., 2007 ). In birds, more aggressive or risk-prone individuals often perform more poorly in tasks requiring response inhibition or motor control, such as detour paradigms (Speechley et al., 2024 ; Vernouillet et al., 2025 ), although evidence remains mixed and appears context- and task-dependent (van Iersel et al., 2025 ; Willcox et al., 2026 ; Willcox, et al., 2025 a). These findings raise the possibility that MA could be associated with reduced inhibitory control. While reduced inhibitory control could contribute to the expression of MA, exposure to MA may also influence the development of inhibitory control in offspring. Adverse early experiences, such as MA or unpredictable parental interactions, have been shown to alter a range of behavioural traits in mammals and birds, including emotional reactivity, stress responsiveness, and anxiety-like behaviours (Anda et al., 2006 ; Arnold & Taborsky, 2010 ; Bannier et al., 2017 ; Gondré-Lewis et al., 2016 ; Lovallo, 2013 ). Executive functions such as inhibitory control are also particularly sensitive to early-life social conditions because they develop during periods of heightened neural plasticity (Bernier et al., 2012 ; Bosquet Enlow et al., 2019 ; Dydenkova et al., 2024 ; McEwen & Morrison, 2013 ; Meaney, 2010 ). Exposure to MA during development may therefore shape offspring's inhibitory control, opening the possibility for transgenerational transmission of MA (Champagne & Meaney, 2007 ; Maestripieri et al., 2007 ). Importantly, because inhibitory control shows only moderate heritability across species (Gnanadesikan et al., 2020 ; Langley et al., 2020 ), genetic transmission alone is unlikely to fully account for such intergenerational continuity, making post-hatching experience a critical pathway to examine. Here, we investigated whether maternal aggression (MA) in the domestic canary ( Serinus canaria) is associated with reduced inhibitory control and whether being exposed to MA as an offspring affects the development of executive functions (i.e., their inhibitory control), which could facilitate an intergenerational transmission of MA. In domestic canaries, females commonly exhibit MA towards their nestlings before fledging (Garcia-Co et al., 2025 ), making it a suitable model to examine potential cognitive processes underlying MA expression and transmission. Inhibitory control was assessed using a detour task, which requires individuals to suppress a direct, prepotent motor response towards a visible reward in favour of an indirect solution (Davidson et al., 2022 ; Kabadayi et al., 2018 ; MacLean et al., 2014 ). Performance was captured by two complementary measures: detour latency, defined as the time taken from entering the test box to circumventing the barrier, which reflects the speed with which a prepotent approach response is inhibited; and persistence, defined as the cumulative time spent approaching the barrier directly without detouring, which reflects the degree to which the prepotent response is repeatedly expressed. If MA reflects deficits in inhibitory control, we predicted that adult females expressing MA would show poorer performance in the detour task test than non-aggressive females, indicated by longer latencies and greater persistence ( prediction 1 ). If early-life exposure to MA shapes offspring inhibitory control, we predicted that offspring raised by aggressive foster mothers would show poorer performance ( prediction 2a) , while offspring born to aggressive biological mothers would perform more poorly if genetic or prenatal maternal effects are involved ( prediction 2b ). We also compared performance between juveniles and adults to characterise ontogenetic differences in inhibitory control and examined whether offspring performance covaried with that of their biological mothers as a measure of heritability. Together, this study aimed to evaluate whether variation in inhibitory control can help explain the expression and potential intergenerational transmission of MA in canaries. MATERIALS AND METHODS STUDY SPECIES AND EXPERIMENTAL SET-UP All birds belonged to a local population of captive Fife fancy canaries kept at the University of Antwerp. To maintain genetic diversity, this population is supplemented annually in late autumn with new individuals obtained from local breeders. In February, all the individuals selected for breeding (approximately 50% locally bred and 50% newly introduced individuals) were housed for five weeks prior to pair formation in single-sex aviaries at a room temperature of 20–24°C with long artificial daylight (14h light: 10h dark; Estramil et al., 2014 ). Breeding pairs were then formed by placing a male followed by a female into a breeding cage (50 x 64 x 40 cm 3 , GEHU cages, The Netherlands). Pairs were initially formed at random, after which relatedness between partners was checked to ensure that no closely related individuals were paired. A total of 103 breeding pairs was established. Each breeding cage was equipped with two perches, a nest cup with nesting material, cuttlefish bone, shell sand, canary seed mixture (Van Camp, Belgium), egg food (provided twice a week; Van Camp, Belgium), and water ad libitum . Nest building was monitored daily, and freshly laid eggs were weighed each morning. Cross-fostering design We cross-fostered entire clutches (n = 103) two to three days after clutch completion, depending on the availability of suitable foster nests. Foster nests were selected based on mean egg mass (< 0.2 g difference), clutch size (± 1 egg), and laying date of the last egg (± 1 day). Cross-fostering was performed blindly with respect to subsequent MA, as this maternal behaviour could only be assessed later in the breeding season. MA phenotypes could be determined for at least one of the two mothers (biological or foster) in 83 nests. However, complete phenotype information for both mothers (i.e., biological and foster mothers) was available for 59 cross-fostered pairs. Missing phenotype information resulted from adult mortality, breeding failure (e.g., failed laying, failed hatching), or complete brood loss before day 14, the developmental stage at which MA is typically expressed. These 59 clutches were retrospectively assigned to four experimental groups, based on whether the MA phenotype of the biological and foster mothers: biological mother that exerted MA – foster mother that did not exert MA (MA-noMA; n = 12); biological mother that exerted MA – foster mother that exerted MA (MA-MA; n = 16); biological mother that did not exert MA – foster mother that did not exert MA (noMA – noMA; n = 17); and biological mother that did not exert MA – foster mother that exerted MA (noMA – MA; n = 14). This cross-fostering design allowed us to disentangle effects of biological origin (including genetic and prenatal maternal influences) from post-hatching rearing environment. Throughout the manuscript, these components are referred to as “biological” and “foster” mother effects, respectively, acknowledging that biological effects may include both genetic inheritance and prenatal influences such as hormone deposition into the egg (Groothuis et al., 2005 ; Vergauwen et al., 2014 ). Nestling period and post-fledging housing From 14 days after laying of the first egg of the clutch onward, nests were checked daily for hatching. Newly hatched nestlings were marked with a non-toxic coloured marker for individual identification. Parents were provided daily with egg food supplemented with freshly germinated seeds throughout the nestling period. Nestlings were weighed every other day with an electronic balance until fledging (± 25 days old). Once nestlings reached 7g, they were fitted with numbered plastic leg rings for permanent identification. Molecular sexing was conducted using blood samples collected at approximately 25 days of age. At fledging (late April to early May), juveniles were separated from their parents and housed in mixed-sex aviaries (2 x 2 x 2m 3 ). Temperature conditions were kept constant (22–24°C), and the light was gradually adjusted to follow the natural seasonal decrease (in one-hour increments starting from a 15h light-9h dark cycle and ending with a 10h light-14h dark cycle). Aviaries were equipped with wooden perches, shell sand, and ad libitum access to canary seed mixture and water. Egg food was provided twice a week. Juveniles were tested at seven weeks of age (approximately three weeks after fledging) and subsequently returned to the population. Categorisation of Exposure to Maternal Aggression Although both parents contribute to parental care post-hatching, male aggression toward offspring is negligible in frequency and intensity (Garcia-Co et al., 2025 ). Maternal behaviour, therefore, represents the main source of early adverse social experience. Males, however, contribute genetically and epigenetically, and could still shape offspring phenotypes, an aspect that deserves future study. Maternal aggression (MA) was defined as feather plucking directed toward at least one nestling in a brood (Garcia-Co et al., 2025 ). Based on this nest-level criterion, clutches were retrospectively assigned to the four experimental groups described above. Of the 70 females for which maternal behaviour could be reliably assessed, 35 exhibited MA and 35 did not. MA typically began between 14 and 19 days after hatching. However, because not all nestlings within a brood with an aggressive mother were necessarily targeted, we additionally recorded individual-level exposure, defined as whether an individual chick was directly plucked by the foster mother. For the offspring, this individual-level measure was used in all statistical analyses as the measure of early-life social stress. In contrast, the biological mother’s MA phenotype was treated as a nest-level trait, defined by whether she displayed MA toward at least one chick in her foster brood during the same breeding season. DETOUR TASK: Inhibitory control was quantified using a detour task that followed exactly the protocol described for canaries in Dewulf et al. ( 2025 ). The task required individuals to inhibit their response to approach a visible food reward directly through a semi-transparent barrier (as this would result in collision with the obstacle) and instead obtain the reward by detouring around either side of the barrier. Both mothers and offspring were tested using this protocol. Mothers were tested before the breeding season to assess whether inhibitory control is associated with the expression of MA. Offspring were tested during the juvenile stage to evaluate whether early-life exposure to maternal aggression predicts variation in inhibitory control. A total of 122 adult females were tested. Eight individuals were excluded based on pre-registered exclusion criteria (three failed habituation; five failed to meet participation criteria during the first test trial; see below “Data exclusion criteria” section for more information). MA phenotype information was available for 70 of the successfully tested females (35 MA and 35 noMA). For offspring, 114 individuals were tested for which both biological and foster mother MA phenotypes were known, and which passed all exclusion criteria. Juveniles were habituated at 41–43 days old and tested at 44–45 days old, corresponding to approximately three weeks post-fledging. The study employed a semi‑transparent barrier composed of either horizontal or vertical bars, following the design used by Dewulf et al. ( 2025 ), for which the juvenile data were originally collected. Barrier type (horizontal vs. vertical bars) and barrier order (which barrier type was presented first) were excluded from the present analyses for simplicity, as no significant effects were detected. Models including barrier type and order are available upon request. Apparatus and set-up The apparatus consisted of a dark, two-door start box connected to a test box containing a floor-to-ceiling transparent Perspex barrier and a coloured food bowl placed directly behind it. For canaries, a standard orange-brown bowl (Elho, Belgium) was used, containing a visible yet restricted food reward (seed mix and egg food). The barrier was made of transparent Perspex with 18 horizontal black lines painted across its surface, scaled according to canary morphology so that the occluding lines masked approximately 14% of the visual area, as in the original study (Zucca et al., 2005 ). Barrier dimensions were validated in other species (see Dewulf et al., 2025 ) and rescaled for use in canaries using tarsus length measured at day 25, which corresponds to the plateau phase of canary morphological development. Floor-to-ceiling barriers were used to prevent birds from flying over the barrier. All trials were recorded using a Sony Action Cam HDR-AS50 mounted centrally above the test box. Habituation Before testing, all birds underwent 10 days of habituation in their home enclosure, during which they were fed from a coloured food bowl identical to the bowl used in the task. Food was provided ad libitum , except on the evening before each habituation or test day, when feeders were removed at 18:00, creating a natural overnight non-feeding period, commonly used in behavioural studies of canaries. Habituation inside the test box consisted of three consecutive days, in which each individual received one trial per day. Birds were placed in the start box and allowed to enter the test box freely. On the second and third habituation days, an opaque barrier was placed immediately behind the food bowl. This configuration allowed us to obtain a multi-baseline measure of each individual’s general motivational state. Because the barrier was opaque and positioned directly behind the food bowl, individuals were required to approach and feed without engaging in a detour or inhibiting a prepotent response. Latencies measured in this context, therefore, reflect a set of factors such as food motivation, test-box neophobia, barrier neophobia, and general exploratory tendency. For each individual, this multi-baseline measure was calculated as the mean latency (in seconds; s) between leaving the start box and touching the food bowl across habituation trials 2 and 3. This measure was subsequently included as a covariate in all statistical models of detour task performance. Habituation trials ended once the bird had fed for 30 s or after 5 min 15 s for juveniles and 2 min 15 s for adults (the longer duration for juveniles was used to accommodate higher neophobia and slower familiarisation typical of recently fledged birds; Greenberg, 2003 ; Higgins et al., 2022 ). The longer habituation trial duration allowed for adequate familiarisation with the apparatus. Testing Testing took place over two consecutive days, with one test session per day. Each test session consisted of three trials using one of the two barrier types (horizontal-barred or vertical-barred barrier), with barrier order pseudo-randomised across individuals. At the start of every trial, the bird was placed inside the dark start box. After 15 s, the opaque front door was opened, allowing the bird to view the test box without entering. After another 15 s, the transparent door was opened, allowing the bird to enter the test box. Birds failing to exit within 30 s were gently encouraged forward by sliding the rear panel of the start box forward. Test trials ended immediately upon food contact or after 2 min 15 s (both for juveniles and adults). The shorter test trial duration prevented birds from becoming satiated or disengaged. Behavioural annotation and variable extraction All behaviours from the second and third habituation trials and from all six test trials were coded using BORIS (v7.13.6; Friard & Gamba, 2016 ). Coding followed the definitions of Dewulf et al., ( 2025 ). For each trial we recorded (i) the latency to leave the start box, (ii) the latency to detour, defined as the interval between leaving the start box and circumventing the barrier, (iii) the time spent persisting at the barrier (i.e. cumulative time inside the barrier zone of interest (see Dewulf et al., 2025 ) and (iv) the moment of food (bowl) contact. Trials in which the bird entered the barrier zone of interest but failed to detour were assigned the maximum detour latency (135 s). Trials in which the bird immediately detoured without entering the zone were assigned zero seconds of persistence. A multi-baseline motivational measure was calculated as the mean latency to reach the food bowl across habituation trials 2 and 3 (or from trial 3 alone when contact was not made in trial 2). The main dependent variables were latency to detour, persistence time, and improvement across trials and test days. Data exclusion criteria Pre-registered exclusion criteria from Dewulf et al., ( 2025 ) were applied. Birds that did not approach the food bowl by the third habituation day were excluded from the testing phase to ensure adequate task proficiency. During testing, birds that neither detoured nor entered the barrier zone of interest were excluded from further trials, as such behaviour likely indicated demotivation or distress. Individuals that left the test box and detoured without contacting the food bowl were also excluded, ensuring that exploratory behaviour was not misinterpreted as successful detouring. Eight individuals were excluded based on pre-registered exclusion criteria (three failed habituation; five failed to meet participation criteria during the first test trial). STATISTICAL ANALYSIS All statistical analyses were performed in R 4.1.3 (R Core Team, 2013 ). For all statistical tests, the significance level was set at α = 0.05. Model assumptions were assessed visually using residual diagnostics generated with the packages performance (v0.10.3; Lüdecke et al., 2021 ) and DHARMa (v0.4.6; Hartig, 2016 ). We used Tukey’s method based on the interquartile range (IQR) and visually inspected scatterplots and residual distributions. When necessary, data were winsorised to limit the influence of extreme values (Wilcox, 2012 ). Linear mixed-effects models (LMMs) and generalised mixed-effects models (GLMMs) were fitted using the lme4 package (v1.1-35.5; Bates et al., 2015 ). For LMMs, p-values were obtained using lmerTest (v3.1.3, Kuznetsova et al., 2017 ), which provides Type III ANOVA tables by default. Beta mixed-effects models were fitted using the glmmTMB package (v1.1.12; Brooks et al., 2017 ). Model selection was based on backward elimination of non-significant interaction terms (α = 0.05), guided by changes in the Akaike Information Criterion (AIC). In the main text, we report results from the minimal adequate models; full model outputs are provided in the Supplementary Information. For clarity, we followed the analytical approach used in recent work (Willcox et al., 2026 ) and collapsed across test sessions, modelling Trial as a single factor with six levels (three trials from each session). The full models—initially including Test session (session 1 vs. session 2), Trial number (1, 2, or 3), and their interaction to account for between-session differences and within-session learning—are reported in the Supplementary Information. Relationship between MA phenotype and detour task performance in adult females (prediction 1) To test prediction 1, we analysed detour task performance using mixed-effects models. Detour latency was analysed using a linear mixed-effects model with MA phenotype (MA vs noMA) as a fixed effect. Trial number (1–6) and multi-baseline measure (mean food-approach latency during habituation with an opaque barrier) were included as covariates. Female identity was included as a random intercept to account for repeated measures across trials. Detour latency values were positively skewed; therefore, the response variable was winsorised using a trim value of 0.05 prior to analysis to reduce the influence of extreme values while retaining all observations. Persistence at the barrier was quantified as the proportion of trial time spent persisting and analysed using a beta mixed-effects model with a logit link function. Because trial duration differed between successful and unsuccessful trials in adult females, a proportional measure was required to ensure comparability across observations. Values of 0 and 1 were adjusted following standard recommendations for beta regression Smithson & Verkuilen, ( 2006 ). The same fixed and random effects structure as in the detour latency model was applied. Improvement in detour latency across trials could be inferred from the MA phenotype × Trial interaction term included in the detour latency model described above. Effects of early-life exposure to MA on offspring detour task performance (predictions 2a and 2b) To test predictions 2a and 2b, we analysed offspring detour task performance using mixed-effects models that accounted for the cross-fostering design. The MA phenotype of the biological mother and the MA phenotype of the foster mother were included as separate fixed effects, along with their interaction, allowing us to disentangle biological (genetic and prenatal maternal) effects from post-hatching environmental effects. Detour latency was analysed using a linear mixed-effects model with winsorised latency values (trim value = 0.05). Trial number and multi-baseline measure were included as covariates. Individual identity nested within nest identity was included as a random intercept to account for repeated measures across trials and non-independence among siblings. For offspring, persistence at the barrier was quantified as the total time spent persisting, rather than as a proportion. Because all offspring test trials were of identical duration, absolute persistence times were directly comparable across individuals and trials. Persistence was analysed using a linear mixed-effects model with the same fixed and random effects structure as for detour latency. Improvement in detour latency across trials could be inferred from the interactions of biological and foster mother MA phenotype with Trial included in the main latency model described above. Additional analyses a) Age-related differences in detour task performance Because inhibitory control develops during ontogeny, we compared juveniles and adults using mixed-effects models, including age class (juvenile vs adult) as a fixed effect. Trial number was included as a covariate. Individual identity was included as a random intercept to account for repeated measures. Detour latency and persistence were analysed as described above. Improvement across trials could be inferred from the Age class × Trial interaction included in both models. b) Mother-offspring resemblance in detour task performance To provide exploratory insight into potential intergenerational similarity in inhibitory control, we examined whether offspring detour task performance covaried with the performance of their biological mothers using a mid-offspring regression approach. For each trait, detour latency, mean persistence (proportion of time spent persisting), and improvement across trials, we first calculated a single value per individual by averaging across test sessions. Offspring values were then averaged at the nest level to obtain one mean offspring value per family. These nest-level offspring means were paired with the corresponding maternal values, resulting in one data point per family. This approach avoids pseudo-replication arising from multiple offspring per mother and provides an estimate of intergenerational resemblance at the family level. We fitted linear models in which mean offspring performance was used as the response variable and maternal performance as a continuous predictor. For detour latency, both maternal and offspring values were log-transformed prior to analysis. Persistence values were analysed on the original scale. Improvement was calculated as the difference in detour latency between Trial 1 and Trial 6 (Trial 1 − Trial 6), such that positive values indicate improved performance across trials. Offspring improvement scores were square-root transformed to meet model assumptions, whereas maternal improvement was entered on the raw scale. RESULTS Relationship between MA phenotype and detour task performance in adult females (prediction 1) The MA phenotype was not associated with detour task performance. Adult females who later expressed MA did not differ from non-aggressive females in their detour latency ( MA : p = 0.38; Table 1 ; Fig. 1 a). Across all adult females, detour latency decreased significantly across trials ( Trial : p < 0.001), indicating learning over repeated exposures to the task. Adult females who later expressed MA did not spend a different proportion of time persisting at the barrier than females that did not express MA ( MA : p = 0.39; Table 1 ; Fig. 1 b). No trial effect was detected for the proportion of time persisting ( Trial : p = 0.16). Multi-baseline measure (i.e., mean food-approach latency during habituation with an opaque barrier; see Materials and methods: Habituation ) did not significantly explain variation in either detour latency or proportion of time persisting (Table 1 ). Table 1 Outcome of models testing the association between maternal aggression (MA) phenotype and detour task performance in adult females. Proportion of time persisting was adjusted to accommodate values of 0 and 1. Bold values indicate P < 0.05. Detour Latency Estimate (SE) F df P Intercept 2.06 (± 0.15) - - - MA -0.14 (± 0.16) 0.77 (1, 42.45) 0.38 Multi-baseline measure -0.01 (± 0.01) 0.15 (1, 46.42) 0.69 Trial -0.20 (± 0.02) 204.81 (1, 204.81) < 0.001 Estimate (SE) Z value P Proportion of time persisting Intercept 0.28 (± 0.18) 1.53 0.12 MA -0.14 (± 0.17) -0.85 0.39 Multi-baseline measure 0.01 (± 0.01) 0.28 0.77 Trial -0.05 (± 0.03) -1.38 0.16 Effects of early-life exposure to MA on offspring detour task performance (prediction 2a and 2b) There was no evidence that early-life exposure to MA was associated with offspring detour task performance. Neither post-hatching exposure to MA ( foster mother phenotype : p = 0.59) nor the MA phenotype of the biological mother ( biological mother phenotype : p = 0.13; Table 2 ; Fig. 2 a) significantly explained variation in offspring detour latency. Similarly, total time spent persisting at the barrier did not differ according to foster ( foster mother phenotype : p = 0.73) or biological ( biological mother phenotype : p = 0.89) mother MA phenotype (Table 3 ; Fig. 2 b). Across offspring, detour latency and total time persisting both decreased significantly across trials ( Trial : p < 0.001 in both cases), indicating learning over repeated exposures to the task. The multi-baseline measure was a significant covariate for detour latency (p = 0.04; Table 2 ): individuals with longer food-approach latencies during habituation also showing longer detour latencies during testing. It did not significantly predict total time persisting (see Table 2 ). Table 2 Outcome of mixed-effects models testing the relationship between maternal aggression (MA) exposure and detour task performance in offspring. Detour latency and total time persisting were analysed using linear mixed-effects models. Detour latency was winsorised (trim = 0.05). Groups were defined by the MA phenotype of the biological and foster mothers. Bold values indicate P < 0.05. Detour latency Estimate (SE) F df P Intercept 2.96 (± 0.12) - - < 0.001 Multi-baseline measure 0.01 (± 0.01) 4.04 (1, 105.72) 0.04 Biological mother phenotype -0.22 (± 0.14) 2.24 (1, 108.59) 0.13 Foster mother phenotype 0.08 (± 0.15) 0.27 (1, 111.73) 0.59 Trial -0.27 (± 0.01) 306.92 (1, 566.51) < 0.001 Estimate (SE) F df P Total time persisting Intercept 8.80 (± 0.85) - - < 0.001 Multi-baseline measure 0.01 (± 0.01) 1.19 (1, 94.81) 0.27 Biological mother phenotype 0.11 (± 0.93) 0.01 (1, 89.30) 0.89 Foster mother phenotype -0.33 (± 0.98) 0.11 (1, 108.05) 0.73 Trial -1.64 (± 0.17) 91.33 (1, 571.10) < 0.001 Age-related differences in detour task performance Detour task performance differed markedly between juveniles and adults. For detour latency, there was a strong main effect of age class (p < 0.001; Table 3 ; Fig. 3 a), with juveniles showing longer detour latencies than adults. Latency decreased significantly across trials (p < 0.001), indicating overall improvement in task performance. Importantly, the interaction between age class and trial number was significant (p = 0.04), showing that the rate of improvement differed between age classes, with juveniles exhibiting a steeper decline in latency across trials than adults. A similar pattern emerged for persistence at the barrier. There was a significant effect of age class (p < 0.001; Table 4; Fig. 3 b), with juveniles spending less time persisting at the barrier than adults. Persistence decreased across trials (p = 0.02), consistent with improved task performance over time. The interaction between age class and trial was also significant (p = 0.02), indicating that juveniles showed a steeper reduction in persistence across trials, while adult persistence remained comparatively stable, consistent with the absence of a significant trial effect for adult persistence observed in the prediction 1 analyses ( Trial : p = 0.16; see Table 1 ). Table 3 Outcome of models testing the association between age class and detour task performance. Detour latency and proportion of time persisting were analysed using linear mixed-effects models. Detour latency was log-transformed and winsorised (trim = 0.05). Proportion of time persisting at the barrier was square-root transformed. Bold values indicate P < 0.05. Detour latency Estimate (SE) F df P Intercept 2.13 ( ± 0.09) - - < 0.001 Age class 0.82 ( ± 0.12) 41.79 (1, 511.22) < 0.001 Trial -0.22 ( ± 0.01) 134.25 (1, 985.90) < 0.001 Age class: Trial -0.04 ( ± 0.02) 4.01 (1, 963.89) 0.04 Estimate (SE) F df P Proportion of time persisting Intercept 1.16 ( ± 0.06) - - < 0.001 Age class -0.61 ( ± 0.11) 31.34 (1, 170.98) < 0.001 Trial -0.04 ( ± 0.01) 4.89 (1, 268.32) 0.02 Age class: Trial 0.09 ( ± 0.04) 4.79 (1, 282.24) 0.02 Mother-offspring resemblance in detour task performance We found no evidence for mother-offspring resemblance in inhibitory control. Mean offspring detour latency was not associated with maternal detour latency (β = −0.12 ± 0.14 SE, F ₁, ₂₉ = 0.75, p = 0.39; Fig. 4 a), indicating that offspring from mothers with slower or faster detour performance did not differ in their own latency to solve the task. Similarly, offspring persistence at the barrier was not related to maternal persistence (β = −0.05 ± 0.08 SE, F ₁, ₂₉ = 0.41, p = 0.53; Fig. 4 b), providing no evidence for intergenerational similarity in this behavioural component of the task. Consistent with these findings, improvement in detour performance across trials was not associated with maternal improvement (β = -0.08 ± 0.07, F ₁, 22.63 = 1.30, p = 0.26; Fig. 4 c), indicating that offspring did not resemble their mothers in their rate of performance change over repeated trials. DISCUSSION This study examined whether maternal aggression (MA) is associated with individual differences in inhibitory control in adult canary females and whether early-life exposure to MA contributes to variation in offspring inhibitory control. Contrary to our predictions, we found no evidence that MA phenotype in adult females was associated with detour task performance, indicating that aggressive and non-aggressive mothers did not differ in their ability to inhibit prepotent motor responses. Despite representing a harsh early-life social environment, post-hatching exposure to MA did not influence offspring inhibitory control. This absence of environmental effects is notable given that executive functions are often considered sensitive to early developmental conditions. Detour performance varied with age: juveniles showed longer detour latencies than adults and improved more steeply across trials, suggesting that behavioural performance in this task continues to develop after hatching. However, adults spent more time persisting at the barrier than juveniles. Finally, we found no evidence for mother-offspring resemblance in detour task performance. Maternal aggression is not associated with impaired inhibitory control in adult females The absence of an association between MA and detour task performance in adult females suggests that MA in canaries is unlikely to reflect reduced response inhibition or heightened impulsivity, at least as captured by this task. While some studies have reported links between aggression and inhibitory control in birds, most notably in Australian magpies ( Ghymnorhina tibicen ), where aggressive or risk-prone individuals performed more poorly in tasks requiring motor control or behavioural flexibility (Speechley et al., 2024 ; but see also, Vernouillet et al., 2025 ), our findings align with a growing body of work that fails to replicate such associations. For example, van Iersel et al. ( 2025 ) found no relationship between territorial aggression and detour task performance in free-living female blue tits ( Cyanistes caeruleus ), and Willcox et al., ( 2026 ) similarly reported no association between aggression and response inhibition in chickens ( Gallus gallus domesticus ). Along with our study, these recent findings challenge the assumption of a generalised cognitive-behavioural syndrome linking aggression to impaired inhibition. However, detour tasks typically involve not only response inhibition, but also problem-solving, spatial learning, and exploration, meaning that environment-dependent effects on inhibitory control specifically may be diluted at the task level (Kabadayi et al., 2018 ; van Horik et al., 2018 ). Thus, the measures we obtained are composite, and this may limit their sensitivity to specific regulatory processes. Moreover, performance often shows low consistency across different inhibition tasks, suggesting that task design strongly influences which inhibitory processes are captured (Brucks et al., 2017 ; Troisi et al., 2026a , 2026b ; van Horik et al., 2018 ; Völter et al., 2018 ). This should be borne in mind when interpreting the present study. That said, task-level limitations alone cannot fully account for the absence of an association between MA and detour performance, since they would equally apply to studies that do detect such links between aggression and inhibitory control. A more fundamental explanation may therefore lie in the nature of MA itself. MA in canaries may not represent reactive or impulsive aggression, but rather a context-dependent component of a parental investment strategy. From this perspective, MA may represent an escalation of selective investment decisions rather than deficits in behavioural regulation. Comparable forms of parental aggression or brood reduction have been documented in birds such as coots ( Fulica atra) , storks ( Ciconia ciconia ), and pelicans ( Pelecanus erythrorhynchos ), where aggression towards offspring facilitates brood reduction under limited resources (see Cash & Evans, 1986 ; Shizuka & Lyon, 2013 ; Tortosa & Redondo, 1992 ; Zieliński, 2002 ), as well as in mammals, where maternal rejection or aggression biases investment towards higher-quality offspring (e.g., Maestripieri, 1998 ; Mock & Parker, 1997 ). If MA serves as a strategic adjustment rather than an impulsive response, it would not be expected to co-vary with individual differences in inhibitory control measured in a non-social, non-breeding context. Interestingly, MA females did not score higher than noMA females on a social dominance test conducted outside the breeding context (Garcia-Co et al., in prep.), suggesting that MA does not reflect a broadly aggressive phenotype. Rather, aggressiveness in the breeding context appears to be decoupled from social behaviour in other contexts, consistent with the view that MA is a state-dependent, context-specific trait (Dingemanse et al., 2012 ; Réale et al., 2007 ). Aggression is a multi-faceted behaviour, regulated by distinct neural and endocrine mechanisms depending on its functional context (e.g., territorial, social, or parental), and individuals often express aggression selectively rather than consistently across situations (Dingemanse et al., 2012 ; Maney & Goodson, 2011 ; Nelson & Trainor, 2007 ; Yanowitch & Coccaro, 2011 ). If MA is not part of a broader, cross-context aggressive phenotype, then it is unlikely to be associated with individual differences in inhibitory control measured in a non-social, non-reproductive setting. Together, these considerations suggest that the absence of an association between MA and detour performance reflects the context-specificity of MA expression, rather than a genuine absence of any link between inhibitory control and aggressive behaviour in this species. Early-life exposure to maternal aggression does not alter offspring inhibitory control We found no evidence that offspring raised by aggressive mothers differed in detour task performance from offspring associated with non-aggressive mothers. This absence of post-hatching environmental effects is notable given that MA constitutes an early-life social stressor and has been linked to long-term behavioural differences in this system, including heightened aggressiveness and reduced neophobia (Garcia-Co et al., 2025 , 2026 ). Furthermore, a substantial body of work shows that early-life conditions can shape inhibitory control and related executive functions across taxa (Brett et al., 2015 ; Dydenkova et al., 2024 ; Hedges & Woon, 2011 ; Rowell et al., 2021 ). In birds, for example, experimental manipulations of early life environmental predictability, enrichment, and social conditions have been shown to influence inhibitory control and persistence in detour or cylinder tasks (Ashton et al., 2018 ; van Horik et al., 2019 ; Vernouillet et al., 2025 ; Willcox et al., 2025 a), although such effects are not consistently detected (Willcox et al., 2026 ). Several considerations may explain this discrepancy. First, as noted above, detour performance is a composite measure while early-life MA may selectively affect inhibitory processes without this being detectable at the level of overall detour performance. Indeed, in a previous study examining responses to an escape test (Garcia-Co et al., 2026 ), offspring born to biological mothers that later exerted MA spent more time pecking at a closed barrier, a measure of persistence, than nestlings born to non-aggressive biological mothers. This suggests that early-life MA does shape some aspects of behavioural regulation, but that these effects may be domain-specific and not generalise across task contexts. Effects may also be more apparent in socially embedded tasks where the inhibitory demands most relevant to an adverse early social environment are more directly engaged. Consistent with this, Willcox et al., ( 2026 ) found that early-life social instability in chickens affected aggression in a social context but not response inhibition in a non-social context. Second, we observed strong age-related differences in detour task performance, with juveniles showing longer detour latencies than adults but improving more steeply over trials, consistent with a gradual maturation of executive functions. The steeper improvement in juveniles likely reflects greater room for improvement at this developmental stage rather than higher learning capacity per se, as individuals starting from a lower baseline have more potential for short-term gain. Comparable ontogenetic trajectories are well documented across birds and mammals, with inhibitory control improving as neural circuits mature and individuals accumulate experience (Diamond, 1990 ; Diamond & Gilbert, 1989 ; Kabadayi et al., 2017 ). Canaries are an altricial species, meaning that neural development continues well after hatching (Iwaniuk & Nelson, 2003 ). Our juveniles were tested shortly after independence, at a stage when inhibitory circuits may not have been fully established at the juvenile stage (Diamond, 1990 ; Kabadayi et al., 2017 ). If the neural systems underlying detour performance are still maturing at the time of testing, they may also be less responsive to the influence of earlier social experience, potentially masking effects of early-life MA that might only emerge later in development. Indeed, in some species the effects of early social environment on cognitive performance have been detected only later in ontogeny the effects of early social environment on cognitive performance emerged only later in ontogeny (Ashton et al., 2018 ; but see van Horik et al., 2019 ; Vernouillet et al., 2025 ; Willcox, et al., 2025 ). Whether MA-exposed offspring would show differences in inhibitory control once fully mature remains an open question. Interestingly, the pattern for persistence was the reverse: adults spent more time persisting at the barrier than juveniles throughout testing. This could reflect greater goal-directedness or motivation in adults, as mature individuals may invest more effort in a direct approach before switching strategies, whereas juveniles might disengage more rapidly after an unsuccessful attempt. Notably, despite their higher persistence, adults still showed generally fast detour latencies overall, suggesting that the time spent near the barrier did not substantially delay task success. This may indicate that repeated barrier approaches carry a low behavioural cost for adults in this context (e.g., Kabadayi et al., 2018 ; van Horik et al., 2018 ), and that adult persistence does not necessarily reflect an inhibitory deficit but rather a different pattern of task exploration. Furthermore, juveniles showed a steeper reduction in persistence across trials than adults, mirroring the steeper improvement in detour latency. Whether this reflects greater neural plasticity or simply lower initial motor fixation on the barrier remains to be determined. Regardless of the underlying mechanism, these results suggest that while inhibitory control improves with age, other motivational factors, such as persistence, may follow a distinct developmental trajectory. We found no evidence for mother-offspring resemblance in detour task performance, suggesting limited genetic or prenatal maternal effects in this measure of inhibitory control. This is consistent with low heritability estimates reported for inhibitory control and related cognitive traits in birds (Gnanadesikan et al., 2020 ; Langley et al., 2020 ). Together with the lack of early environmental effects, the absence of genetic effects supports the view that inhibitory control measures are among others shaped by an individual's experience and current motivational state (Bateson et al., 2015 ; Dunn et al., 2019a ). Limited resemblance may additionally reflect the high plasticity of inhibitory control as a trait. Measures that are strongly responsive to individual experience and current context are inherently more difficult to establish as heritable from a limited number of observations per individual (Brucks et al., 2017 ; McCallum & Shaw, 2023 ; Troisi et al., 2021 ). Conclusions Overall, our results suggest that MA in canaries is not associated with a general impairment in inhibitory control and does not appear to be transmitted across generations through variation in this cognitive trait. The absence of an association between MA and detour performance, together with the lack of cross-context consistency in aggressive behaviour, suggests that MA is a state-dependent, context-specific maternal behaviour rather than the expression of a stable inhibitory deficit. Evidence from a previous study in this population (Garcia-Co et al., 2026 ) nevertheless suggests that early-life MA does selectively shape some aspects of behavioural regulation, pointing to domain-specific effects. Declarations DATA AVAILABILITY The data used in this study is archived in Mendeley Data in the following link: https://data.mendeley.com/datasets/cy8fhzsy56/1. The videos analysed are available from the corresponding author (Clara Garcia-Co) upon request. AUTHOR CONTRIBUTIONS Clara Garcia-Co: Data curation; Formal analysis; Investigation; Methodology; Validation; Visualization; Writing – original draft & editing. Anneleen Dewulf: Data curation; Formal analysis; Investigation; Methodology; Validation; Writing – review & editing. Judith Morales: Funding acquisition; Supervision; Validation; Writing – review & editing. Frederick Verbruggen: Funding acquisition; Supervision; Validation; Writing – review & editing. Wendt Müller: Conceptualization; Funding acquisition; Investigation (supporting); Methodology (supporting); Resources; Supervision; Validation; Writing – review & editing. CONFLICT OF INTEREST STATEMENT The authors declare no competing or financial interests. ACKNOWLEDGEMENTS We thank Peter Scheys for his assistance in taking care of the birds. FUNDING STATEMENT This work was supported by the FWO Flanders (CG: 1173825N and AD: 11F0823N). FV was supported by an ERC Consolidator grant (European Union’s Horizon 2020 research and innovation programme, grant agreement No 769595) and Methusalem Project 01M00221 (Ghent University). JM was supported by PID2022-139166NB-I00 funded by MCIN/AEI/https://doi.org/10.13039/501100011033 and “ERDF A way of making Europe”. STATEMENT OF ANIMAL ETHICS The experiments described in this study were carried out with the approval of the Ethical Committee for Animal Experimentation of the University of Antwerp (ECD 2022-87) and by the Ethical Committee for Animal Experimentation, Faculty of Sciences, Flemish Institute for Biotechnology (VIB) of Ghent University (EC2022-091). We complied with Belgian and Flemish animal welfare legislation, adhered to the ASAB/ABS guidelines for the use of animals in behavioural research and teaching, and ARRIVE guidelines. A constant effort was made to keep the birds in semi-natural conditions and in good health. The invasiveness of all observational procedures described was minimized, and handling time was kept under two minutes per bird. Birds were handled exclusively by trained personnel (AD, CG, WM: FELASA C; caretakers: FELASA B). Body mass and behaviour were monitored throughout the whole experiment, as they reflect general well-being (see Paul-Murphy & Hawkins, 2015). Nestlings were temporarily marked with non-toxic coloured markers until they reached approximately 7 g, when they were fitted with standard metal leg rings; these methods have negligible effects in passerines (Eldegard et al., 2024). For short-term visual identification in group tests, a very small (< 2 mm) dot of correction fluid was applied to crown feathers only. This method is consistent with field marking guidance and produces no observable irritation (Department of Biodiversity, Conservation and Attractions, 2020; NORECOPA, 2022). For food-motivated tasks, birds were food-restricted overnight (for a maximum of 18 hours) to standardise motivation across individuals; this mild deprivation is within natural foraging intervals (see e.g., Cauchoix et al., 2017; Dunn et al., 2019a; Vergauwen et al., 2014). 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Anim Cogn. https://doi.org/10.1007/s10071-025-02033-1 van Horik JO, Beardsworth CE, Laker PR, Langley EJG, Whiteside MA, Madden JR (2019) Unpredictable environments enhance inhibitory control in pheasants. Anim Cogn 22(6):1105–1114. https://doi.org/10.1007/s10071-019-01302-0 van Horik JO, Langley EJG, Whiteside MA, Laker PR, Beardsworth CE, Madden JR (2018) Do detour tasks provide accurate assays of inhibitory control? Proceedings of the Royal Society B: Biological Sciences , 285 (1875), 20180150. https://doi.org/10.1098/rspb.2018.0150 van Iersel R, Pinxten R, Eens M (2025) Is aggression related to inhibitory control in a free-living female songbird? Behav Ecol 36(6):araf126. https://doi.org/10.1093/beheco/araf126 Veenema AH (2009) Early life stress, the development of aggression and neuroendocrine and neurobiological correlates: What can we learn from animal models? Front Neuroendocr 30(4):497–518. https://doi.org/10.1016/j.yfrne.2009.03.003 Verbruggen F, Logan GD (2008) Response inhibition in the stop-signal paradigm. Trends Cogn Sci 12(11):418–424. https://doi.org/10.1016/j.tics.2008.07.005 Vergauwen J, Groothuis TGG, Eens M, Müller W (2014) Testosterone influences song behaviour and social dominance – But independent of prenatal yolk testosterone exposure. Gen Comp Endocrinol 195:80–87. https://doi.org/10.1016/j.ygcen.2013.10.014 Vernouillet A, Willcox K, Allaert R, Dewulf A, Zhang W, Troisi C, Knoch S, Martel A, Lens L, Verbruggen F (2025) To peck or not to peck: The influence of early-life social environment on response inhibition and impulsive aggression in Japanese quails. Royal Society Open Science , 12 . https://doi.org/10.1098/rsos.242228 Völter CJ, Tinklenberg B, Call J, Seed AM (2018) Comparative psychometrics: Establishing what differs is central to understanding what evolves. Philosophical Trans Royal Soc B: Biol Sci 373(1756):20170283. https://doi.org/10.1098/rstb.2017.0283 Wilcox RR (2012) Introduction to Robust Estimation and Hypothesis Testing . Academic Press. ScienceDirect. http://www.sciencedirect.com:5070/book/monograph/9780123869838/introduction-to-robust-estimation-and-hypothesis-testing?via=ihub%3D Willcox K, Vernouillet A, Lens L, Verbruggen F (2025) Early-life group size influences response inhibition, but not the learning of it, in Japanese quails. Learn Behav 53(2):157–170. https://doi.org/10.3758/s13420-024-00643-2 Willcox K, Vernouillet A, Szabo B, Martel A, Lens L, Verbruggen F (2025) Early-life social instability affects aggression, but not response inhibition, in chickens (p. 2025.10.29.685273). bioRxiv. https://doi.org/10.1101/2025.10.29.685273 Willcox K, Vernouillet A, Szabo B, Martel A, Lens L, Verbruggen F (2026) Early-life social instability affects aggression, but not response inhibition, in chickens. Acta Ethologica 29(1):5. https://doi.org/10.1007/s10211-025-00477-9 Yanowitch R, Coccaro EF (2011) The Neurochemistry of Human Aggression. In Advances in Genetics (Vol. 75, pp. 151–169). Academic Press. https://doi.org/10.1016/B978-0-12-380858-5.00005-8 Zhang Z, Wang Q, Liu X, Song P, Yang B (2017) Differences in Inhibitory Control between Impulsive and Premeditated Aggression in Juvenile Inmates. Frontiers in Human Neuroscience , 11 . https://doi.org/10.3389/fnhum.2017.00373 Zieliński P (2002) Brood Reduction and Parental Infanticide—Are the White Stork Ciconia ciconia and the Black Stork C. nigra exceptional? Acta Ornithologica 37(2):113–119. https://doi.org/10.3161/068.037.0207 Zucca P, Antonelli F, Vallortigara G (2005) Detour behaviour in three species of birds: Quails (Coturnix sp.), herring gulls (Larus cachinnans) and canaries (Serinus canaria). Anim Cogn 8(2):122–128. https://doi.org/10.1007/s10071-004-0243-x Additional Declarations No competing interests reported. Supplementary Files SIMS4forsubmission.docx Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 12 May, 2026 Reviewers invited by journal 20 Apr, 2026 Editor assigned by journal 15 Apr, 2026 Submission checks completed at journal 15 Apr, 2026 First submitted to journal 13 Apr, 2026 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-9402003","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":626632114,"identity":"daaa666d-c4d5-4af9-bc3d-7e941d2da3bd","order_by":0,"name":"Clara Garcia-Co","email":"data:image/png;base64,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","orcid":"","institution":"University of Antwerp","correspondingAuthor":true,"prefix":"","firstName":"Clara","middleName":"","lastName":"Garcia-Co","suffix":""},{"id":626632115,"identity":"6a10227c-99a6-4043-a896-b443e1a2fa70","order_by":1,"name":"Anneleen Dewulf","email":"","orcid":"","institution":"Ghent University","correspondingAuthor":false,"prefix":"","firstName":"Anneleen","middleName":"","lastName":"Dewulf","suffix":""},{"id":626632116,"identity":"2eee6df7-3772-4cbe-81af-9a007aab2d93","order_by":2,"name":"Frederick Verbruggen","email":"","orcid":"","institution":"Ghent University","correspondingAuthor":false,"prefix":"","firstName":"Frederick","middleName":"","lastName":"Verbruggen","suffix":""},{"id":626632117,"identity":"978921d0-6db8-4de4-8040-af28ce194ea3","order_by":3,"name":"Judith Morales","email":"","orcid":"","institution":"National Museum of Natural Sciences, Spanish National Research Council (MNCN-CSIC","correspondingAuthor":false,"prefix":"","firstName":"Judith","middleName":"","lastName":"Morales","suffix":""},{"id":626632118,"identity":"1f614965-9d1a-47c0-9d5c-4b84df48d221","order_by":4,"name":"Wendt Müller","email":"","orcid":"","institution":"University of Antwerp","correspondingAuthor":false,"prefix":"","firstName":"Wendt","middleName":"","lastName":"Müller","suffix":""}],"badges":[],"createdAt":"2026-04-13 09:42:38","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9402003/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9402003/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":108181636,"identity":"84e50faa-5499-4a66-8d37-b98fea4b2897","added_by":"auto","created_at":"2026-04-30 08:58:48","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":436702,"visible":true,"origin":"","legend":"\u003cp\u003eDetour latency and persistence in relation to maternal aggression (MA) in adult females. Shown are mean ± SE \u003cstrong\u003e(a)\u003c/strong\u003edetour latency and \u003cstrong\u003e(b)\u003c/strong\u003e proportion of time spent persisting at the barrier across six trials of the detour task, grouped by MA phenotype of the mother (MA = later expressed maternal aggression; noMA = no maternal aggression). MA females are represented by orange dashed lines and noMA females by purple solid lines. Jittered points are the raw data points per individual.\u003c/p\u003e","description":"","filename":"image1.png","url":"https://assets-eu.researchsquare.com/files/rs-9402003/v1/03242d791b123d8e601c40eb.png"},{"id":108102138,"identity":"170a1039-ce3e-485d-af62-41065c95c332","added_by":"auto","created_at":"2026-04-29 11:03:58","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":646530,"visible":true,"origin":"","legend":"\u003cp\u003eDetour latency and persistence across trials in offspring born to and/or raised by mothers differing in maternal aggression (MA).\u003cstrong\u003e \u003c/strong\u003eShown are individual trial-level \u003cstrong\u003e(a)\u003c/strong\u003e detour latencies and \u003cstrong\u003e(b)\u003c/strong\u003e total time spent persisting at the barrier. Solid lines and error bars represent group means ± SE. Groups are defined by the maternal aggression (MA) phenotype of the biological mother (left term) and the foster mother (right term), where \"noMA\" indicates the mother did not express MA and \"MA\" indicates she did (e.g., \"noMA–MA\" denotes offspring of a noMA biological mother raised by an MA foster mother).\u003c/p\u003e","description":"","filename":"image2.png","url":"https://assets-eu.researchsquare.com/files/rs-9402003/v1/eaf81668924e7a5ae8668ee1.png"},{"id":108181862,"identity":"14c057aa-dd24-4e17-b0b6-51d4e37d1ba8","added_by":"auto","created_at":"2026-04-30 08:58:58","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":524574,"visible":true,"origin":"","legend":"\u003cp\u003eAge-related differences in detour task performance across trials. Shown are mean ± SE \u003cstrong\u003e(a)\u003c/strong\u003e detour latency and \u003cstrong\u003e(b)\u003c/strong\u003eproportion of time spent persisting across six trials for juveniles and adults. Adults are represented by black filled circles and dashed lines; juveniles are represented by grey open circles and solid lines. Semi-transparent points represent individual trial-level observations. In panel (a), the y-axis is restricted to 0–40 s for visual clarity.\u003c/p\u003e","description":"","filename":"image3.png","url":"https://assets-eu.researchsquare.com/files/rs-9402003/v1/72c8e5d8cb713a9d394f954c.png"},{"id":108102140,"identity":"0c5e35d2-59da-4c5a-9094-c6a1821f7c60","added_by":"auto","created_at":"2026-04-29 11:03:58","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":600486,"visible":true,"origin":"","legend":"\u003cp\u003eRelationship between mother and offspring detour task performance.\u003cstrong\u003e \u003c/strong\u003eShown are associations between maternal and offspring \u003cstrong\u003e(a)\u003c/strong\u003e mean detour latency (log-transformed), \u003cstrong\u003e(b) \u003c/strong\u003emean persistence (proportion of time spent persisting), and \u003cstrong\u003e(c) \u003c/strong\u003eimprovement in detour latency from Trial 1 to Trial 6 (square-root transformed). Each point represents a single family (one offspring per mother). Solid lines indicate linear model fits with 95% confidence intervals. Axes reflect transformations applied for visualisation; statistical analyses were conducted on log-transformed latency, and square-root transformed improvement values.\u003c/p\u003e","description":"","filename":"image4.png","url":"https://assets-eu.researchsquare.com/files/rs-9402003/v1/e1f08d945bcc92d1a09e6177.png"},{"id":108184527,"identity":"f157da4c-83bd-4832-a00b-5c430cd801db","added_by":"auto","created_at":"2026-04-30 09:04:21","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2574452,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9402003/v1/2199fc38-bac6-476b-99b1-06453e6fc0c1.pdf"},{"id":108102136,"identity":"1c0b9156-f554-43ba-af91-cd734123e514","added_by":"auto","created_at":"2026-04-29 11:03:58","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":52291,"visible":true,"origin":"","legend":"","description":"","filename":"SIMS4forsubmission.docx","url":"https://assets-eu.researchsquare.com/files/rs-9402003/v1/a4fa0a999d08e7be20fbbbac.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"No evidence for a link between maternal aggression and inhibitory control across generations in domestic canaries","fulltext":[{"header":"INTRODUCTION","content":"\u003cp\u003eParental care is a fundamental component of the offspring's early-life environment and plays a crucial role in shaping cognitive and behavioural development. Even though both sexes may contribute to parental care in most species, often mothers have a greater impact, as they can influence offspring development to a larger extent both prenatally, e.g., through hormone deposition in eggs, and postnatally, through more frequent behavioural interactions during which they provide resources, protection, and social cues (Bagot \u0026amp; Meaney, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Crews, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Groothuis et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Reddon, \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). These maternal effects are increasingly recognised as key drivers of individual differences in behaviour and cognition, which can persist across life stages (Bagot \u0026amp; Meaney, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; McEwen \u0026amp; Morrison, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Meaney, \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Veenema, \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). While maternal care is typically beneficial, maternal behaviour can also include aggressive interactions directed towards the offspring. Maternal aggression (MA), such as pecking, chasing, or attacking offspring, has been documented across taxa and is often expressed under conditions of environmental stress, high competition, or reproductive conflict (Arnold \u0026amp; Taborsky, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Champagne \u0026amp; Meaney, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Garcia-Co et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Leonard et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Maestripieri, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Shizuka \u0026amp; Lyon, \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). When occurring during sensitive periods of offspring development, MA has the potential to shape offspring development too, although likely adversely. This raises important questions about the causation and consequences of MA (Anda et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Arnold \u0026amp; Taborsky, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Bannier et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Gondr\u0026eacute;-Lewis et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Lovallo, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOne potential causal mechanism associated with the expression of MA is impaired inhibitory control. Inhibitory control allows individuals to suppress impulsive or inappropriate actions; it is a core executive function that underpins goal-directed and adaptive behaviours (Diamond, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Verbruggen \u0026amp; Logan, \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). In particular, response inhibition refers to the ability to suppress a prepotent motor response (Aron, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Verbruggen \u0026amp; Logan, \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Impairments in response inhibition have been linked to reactive forms of aggression, which are characterised by rapid, emotionally driven responses to perceived challenges (Berkowitz, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Feilhauer et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Hecht \u0026amp; Latzman, \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Zhang et al., \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Across taxa, reduced response inhibition and heightened impulsivity have been associated to altered functioning of prefrontal circuits involved in behavioural regulation (Madole et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Puiu et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Rudebeck et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). In birds, more aggressive or risk-prone individuals often perform more poorly in tasks requiring response inhibition or motor control, such as detour paradigms (Speechley et al., \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Vernouillet et al., \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), although evidence remains mixed and appears context- and task-dependent (van Iersel et al., \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Willcox et al., \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2026\u003c/span\u003e; Willcox, et al., \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2025\u003c/span\u003ea). These findings raise the possibility that MA could be associated with reduced inhibitory control.\u003c/p\u003e \u003cp\u003eWhile reduced inhibitory control could contribute to the expression of MA, exposure to MA may also influence the development of inhibitory control in offspring. Adverse early experiences, such as MA or unpredictable parental interactions, have been shown to alter a range of behavioural traits in mammals and birds, including emotional reactivity, stress responsiveness, and anxiety-like behaviours (Anda et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Arnold \u0026amp; Taborsky, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Bannier et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Gondr\u0026eacute;-Lewis et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Lovallo, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Executive functions such as inhibitory control are also particularly sensitive to early-life social conditions because they develop during periods of heightened neural plasticity (Bernier et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Bosquet Enlow et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Dydenkova et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; McEwen \u0026amp; Morrison, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Meaney, \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Exposure to MA during development may therefore shape offspring's inhibitory control, opening the possibility for transgenerational transmission of MA (Champagne \u0026amp; Meaney, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Maestripieri et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Importantly, because inhibitory control shows only moderate heritability across species (Gnanadesikan et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Langley et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), genetic transmission alone is unlikely to fully account for such intergenerational continuity, making post-hatching experience a critical pathway to examine.\u003c/p\u003e \u003cp\u003eHere, we investigated whether maternal aggression (MA) in the domestic canary (\u003cem\u003eSerinus canaria)\u003c/em\u003e is associated with reduced inhibitory control and whether being exposed to MA as an offspring affects the development of executive functions (i.e., their inhibitory control), which could facilitate an intergenerational transmission of MA. In domestic canaries, females commonly exhibit MA towards their nestlings before fledging (Garcia-Co et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), making it a suitable model to examine potential cognitive processes underlying MA expression and transmission. Inhibitory control was assessed using a detour task, which requires individuals to suppress a direct, prepotent motor response towards a visible reward in favour of an indirect solution (Davidson et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Kabadayi et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; MacLean et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Performance was captured by two complementary measures: detour latency, defined as the time taken from entering the test box to circumventing the barrier, which reflects the speed with which a prepotent approach response is inhibited; and persistence, defined as the cumulative time spent approaching the barrier directly without detouring, which reflects the degree to which the prepotent response is repeatedly expressed. If MA reflects deficits in inhibitory control, we predicted that adult females expressing MA would show poorer performance in the detour task test than non-aggressive females, indicated by longer latencies and greater persistence (\u003cem\u003eprediction 1\u003c/em\u003e). If early-life exposure to MA shapes offspring inhibitory control, we predicted that offspring raised by aggressive foster mothers would show poorer performance (\u003cem\u003eprediction 2a)\u003c/em\u003e, while offspring born to aggressive biological mothers would perform more poorly if genetic or prenatal maternal effects are involved (\u003cem\u003eprediction 2b\u003c/em\u003e). We also compared performance between juveniles and adults to characterise ontogenetic differences in inhibitory control and examined whether offspring performance covaried with that of their biological mothers as a measure of heritability. Together, this study aimed to evaluate whether variation in inhibitory control can help explain the expression and potential intergenerational transmission of MA in canaries.\u003c/p\u003e"},{"header":"MATERIALS AND METHODS","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eSTUDY SPECIES AND EXPERIMENTAL SET-UP\u003c/h2\u003e \u003cp\u003eAll birds belonged to a local population of captive Fife fancy canaries kept at the University of Antwerp. To maintain genetic diversity, this population is supplemented annually in late autumn with new individuals obtained from local breeders. In February, all the individuals selected for breeding (approximately 50% locally bred and 50% newly introduced individuals) were housed for five weeks prior to pair formation in single-sex aviaries at a room temperature of 20\u0026ndash;24\u0026deg;C with long artificial daylight (14h light: 10h dark; Estramil et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Breeding pairs were then formed by placing a male followed by a female into a breeding cage (50 x 64 x 40 cm\u003csup\u003e3\u003c/sup\u003e, GEHU cages, The Netherlands). Pairs were initially formed at random, after which relatedness between partners was checked to ensure that no closely related individuals were paired. A total of 103 breeding pairs was established. Each breeding cage was equipped with two perches, a nest cup with nesting material, cuttlefish bone, shell sand, canary seed mixture (Van Camp, Belgium), egg food (provided twice a week; Van Camp, Belgium), and water \u003cem\u003ead libitum\u003c/em\u003e. Nest building was monitored daily, and freshly laid eggs were weighed each morning.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eCross-fostering design\u003c/h3\u003e\n\u003cp\u003eWe cross-fostered entire clutches (n\u0026thinsp;=\u0026thinsp;103) two to three days after clutch completion, depending on the availability of suitable foster nests. Foster nests were selected based on mean egg mass (\u0026lt;\u0026thinsp;0.2 g difference), clutch size (\u0026plusmn;\u0026thinsp;1 egg), and laying date of the last egg (\u0026plusmn;\u0026thinsp;1 day). Cross-fostering was performed blindly with respect to subsequent MA, as this maternal behaviour could only be assessed later in the breeding season. MA phenotypes could be determined for at least one of the two mothers (biological or foster) in 83 nests. However, complete phenotype information for both mothers (i.e., biological and foster mothers) was available for 59 cross-fostered pairs. Missing phenotype information resulted from adult mortality, breeding failure (e.g., failed laying, failed hatching), or complete brood loss before day 14, the developmental stage at which MA is typically expressed.\u003c/p\u003e \u003cp\u003eThese 59 clutches were retrospectively assigned to four experimental groups, based on whether the MA phenotype of the biological and foster mothers: biological mother that exerted MA \u0026ndash; foster mother that did not exert MA (MA-noMA; n\u0026thinsp;=\u0026thinsp;12); biological mother that exerted MA \u0026ndash; foster mother that exerted MA (MA-MA; n\u0026thinsp;=\u0026thinsp;16); biological mother that did not exert MA \u0026ndash; foster mother that did not exert MA (noMA \u0026ndash; noMA; n\u0026thinsp;=\u0026thinsp;17); and biological mother that did not exert MA \u0026ndash; foster mother that exerted MA (noMA \u0026ndash; MA; n\u0026thinsp;=\u0026thinsp;14). This cross-fostering design allowed us to disentangle effects of biological origin (including genetic and prenatal maternal influences) from post-hatching rearing environment. Throughout the manuscript, these components are referred to as \u0026ldquo;biological\u0026rdquo; and \u0026ldquo;foster\u0026rdquo; mother effects, respectively, acknowledging that biological effects may include both genetic inheritance and prenatal influences such as hormone deposition into the egg (Groothuis et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Vergauwen et al., \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eNestling period and post-fledging housing\u003c/h3\u003e\n\u003cp\u003eFrom 14 days after laying of the first egg of the clutch onward, nests were checked daily for hatching. Newly hatched nestlings were marked with a non-toxic coloured marker for individual identification. Parents were provided daily with egg food supplemented with freshly germinated seeds throughout the nestling period. Nestlings were weighed every other day with an electronic balance until fledging (\u0026plusmn;\u0026thinsp;25 days old). Once nestlings reached 7g, they were fitted with numbered plastic leg rings for permanent identification. Molecular sexing was conducted using blood samples collected at approximately 25 days of age.\u003c/p\u003e \u003cp\u003eAt fledging (late April to early May), juveniles were separated from their parents and housed in mixed-sex aviaries (2 x 2 x 2m\u003csup\u003e3\u003c/sup\u003e). Temperature conditions were kept constant (22\u0026ndash;24\u0026deg;C), and the light was gradually adjusted to follow the natural seasonal decrease (in one-hour increments starting from a 15h light-9h dark cycle and ending with a 10h light-14h dark cycle). Aviaries were equipped with wooden perches, shell sand, and \u003cem\u003ead libitum\u003c/em\u003e access to canary seed mixture and water. Egg food was provided twice a week. Juveniles were tested at seven weeks of age (approximately three weeks after fledging) and subsequently returned to the population.\u003c/p\u003e\n\u003ch3\u003eCategorisation of Exposure to Maternal Aggression\u003c/h3\u003e\n\u003cp\u003eAlthough both parents contribute to parental care post-hatching, male aggression toward offspring is negligible in frequency and intensity (Garcia-Co et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Maternal behaviour, therefore, represents the main source of early adverse social experience. Males, however, contribute genetically and epigenetically, and could still shape offspring phenotypes, an aspect that deserves future study.\u003c/p\u003e \u003cp\u003eMaternal aggression (MA) was defined as feather plucking directed toward at least one nestling in a brood (Garcia-Co et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Based on this nest-level criterion, clutches were retrospectively assigned to the four experimental groups described above. Of the 70 females for which maternal behaviour could be reliably assessed, 35 exhibited MA and 35 did not. MA typically began between 14 and 19 days after hatching. However, because not all nestlings within a brood with an aggressive mother were necessarily targeted, we additionally recorded individual-level exposure, defined as whether an individual chick was directly plucked by the foster mother. For the offspring, this individual-level measure was used in all statistical analyses as the measure of early-life social stress. In contrast, the biological mother\u0026rsquo;s MA phenotype was treated as a nest-level trait, defined by whether she displayed MA toward at least one chick in her foster brood during the same breeding season.\u003c/p\u003e\n\u003ch3\u003eDETOUR TASK:\u003c/h3\u003e\n\u003cp\u003eInhibitory control was quantified using a detour task that followed exactly the protocol described for canaries in Dewulf et al. (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The task required individuals to inhibit their response to approach a visible food reward directly through a semi-transparent barrier (as this would result in collision with the obstacle) and instead obtain the reward by detouring around either side of the barrier. Both mothers and offspring were tested using this protocol. Mothers were tested before the breeding season to assess whether inhibitory control is associated with the expression of MA. Offspring were tested during the juvenile stage to evaluate whether early-life exposure to maternal aggression predicts variation in inhibitory control.\u003c/p\u003e \u003cp\u003eA total of 122 adult females were tested. Eight individuals were excluded based on pre-registered exclusion criteria (three failed habituation; five failed to meet participation criteria during the first test trial; see below \u0026ldquo;Data exclusion criteria\u0026rdquo; section for more information). MA phenotype information was available for 70 of the successfully tested females (35 MA and 35 noMA). For offspring, 114 individuals were tested for which both biological and foster mother MA phenotypes were known, and which passed all exclusion criteria. Juveniles were habituated at 41\u0026ndash;43 days old and tested at 44\u0026ndash;45 days old, corresponding to approximately three weeks post-fledging.\u003c/p\u003e \u003cp\u003eThe study employed a semi‑transparent barrier composed of either horizontal or vertical bars, following the design used by Dewulf et al. (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), for which the juvenile data were originally collected. Barrier type (horizontal vs. vertical bars) and barrier order (which barrier type was presented first) were excluded from the present analyses for simplicity, as no significant effects were detected. Models including barrier type and order are available upon request.\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eApparatus and set-up\u003c/h2\u003e \u003cp\u003eThe apparatus consisted of a dark, two-door start box connected to a test box containing a floor-to-ceiling transparent Perspex barrier and a coloured food bowl placed directly behind it. For canaries, a standard orange-brown bowl (Elho, Belgium) was used, containing a visible yet restricted food reward (seed mix and egg food). The barrier was made of transparent Perspex with 18 horizontal black lines painted across its surface, scaled according to canary morphology so that the occluding lines masked approximately 14% of the visual area, as in the original study (Zucca et al., \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Barrier dimensions were validated in other species (see Dewulf et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) and rescaled for use in canaries using tarsus length measured at day 25, which corresponds to the plateau phase of canary morphological development. Floor-to-ceiling barriers were used to prevent birds from flying over the barrier. All trials were recorded using a Sony Action Cam HDR-AS50 mounted centrally above the test box.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eHabituation\u003c/h3\u003e\n\u003cp\u003eBefore testing, all birds underwent 10 days of habituation in their home enclosure, during which they were fed from a coloured food bowl identical to the bowl used in the task. Food was provided \u003cem\u003ead libitum\u003c/em\u003e, except on the evening before each habituation or test day, when feeders were removed at 18:00, creating a natural overnight non-feeding period, commonly used in behavioural studies of canaries.\u003c/p\u003e \u003cp\u003eHabituation inside the test box consisted of three consecutive days, in which each individual received one trial per day. Birds were placed in the start box and allowed to enter the test box freely. On the second and third habituation days, an opaque barrier was placed immediately behind the food bowl. This configuration allowed us to obtain a multi-baseline measure of each individual\u0026rsquo;s general motivational state. Because the barrier was opaque and positioned directly behind the food bowl, individuals were required to approach and feed without engaging in a detour or inhibiting a prepotent response. Latencies measured in this context, therefore, reflect a set of factors such as food motivation, test-box neophobia, barrier neophobia, and general exploratory tendency. For each individual, this multi-baseline measure was calculated as the mean latency (in seconds; s) between leaving the start box and touching the food bowl across habituation trials 2 and 3. This measure was subsequently included as a covariate in all statistical models of detour task performance.\u003c/p\u003e \u003cp\u003eHabituation trials ended once the bird had fed for 30 s or after 5 min 15 s for juveniles and 2 min 15 s for adults (the longer duration for juveniles was used to accommodate higher neophobia and slower familiarisation typical of recently fledged birds; Greenberg, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Higgins et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The longer habituation trial duration allowed for adequate familiarisation with the apparatus.\u003c/p\u003e\n\u003ch3\u003eTesting\u003c/h3\u003e\n\u003cp\u003eTesting took place over two consecutive days, with one test session per day. Each test session consisted of three trials using one of the two barrier types (horizontal-barred or vertical-barred barrier), with barrier order pseudo-randomised across individuals. At the start of every trial, the bird was placed inside the dark start box. After 15 s, the opaque front door was opened, allowing the bird to view the test box without entering. After another 15 s, the transparent door was opened, allowing the bird to enter the test box. Birds failing to exit within 30 s were gently encouraged forward by sliding the rear panel of the start box forward.\u003c/p\u003e \u003cp\u003eTest trials ended immediately upon food contact or after 2 min 15 s (both for juveniles and adults). The shorter test trial duration prevented birds from becoming satiated or disengaged.\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eBehavioural annotation and variable extraction\u003c/h2\u003e \u003cp\u003eAll behaviours from the second and third habituation trials and from all six test trials were coded using BORIS (v7.13.6; Friard \u0026amp; Gamba, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Coding followed the definitions of Dewulf et al., (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). For each trial we recorded (i) the latency to leave the start box, (ii) the latency to detour, defined as the interval between leaving the start box and circumventing the barrier, (iii) the time spent persisting at the barrier (i.e. cumulative time inside the barrier zone of interest (see Dewulf et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) and (iv) the moment of food (bowl) contact. Trials in which the bird entered the barrier zone of interest but failed to detour were assigned the maximum detour latency (135 s). Trials in which the bird immediately detoured without entering the zone were assigned zero seconds of persistence. A multi-baseline motivational measure was calculated as the mean latency to reach the food bowl across habituation trials 2 and 3 (or from trial 3 alone when contact was not made in trial 2). The main dependent variables were latency to detour, persistence time, and improvement across trials and test days.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eData exclusion criteria\u003c/h2\u003e \u003cp\u003ePre-registered exclusion criteria from Dewulf et al., (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) were applied. Birds that did not approach the food bowl by the third habituation day were excluded from the testing phase to ensure adequate task proficiency. During testing, birds that neither detoured nor entered the barrier zone of interest were excluded from further trials, as such behaviour likely indicated demotivation or distress. Individuals that left the test box and detoured without contacting the food bowl were also excluded, ensuring that exploratory behaviour was not misinterpreted as successful detouring. Eight individuals were excluded based on pre-registered exclusion criteria (three failed habituation; five failed to meet participation criteria during the first test trial).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eSTATISTICAL ANALYSIS\u003c/h2\u003e \u003cp\u003eAll statistical analyses were performed in R 4.1.3 (R Core Team, \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). For all statistical tests, the significance level was set at \u003cem\u003eα\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.05. Model assumptions were assessed visually using residual diagnostics generated with the packages \u003cem\u003eperformance\u003c/em\u003e (v0.10.3; L\u0026uuml;decke et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and \u003cem\u003eDHARMa\u003c/em\u003e (v0.4.6; Hartig, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). We used Tukey\u0026rsquo;s method based on the interquartile range (IQR) and visually inspected scatterplots and residual distributions. When necessary, data were winsorised to limit the influence of extreme values (Wilcox, \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eLinear mixed-effects models (LMMs) and generalised mixed-effects models (GLMMs) were fitted using the lme4 package (v1.1-35.5; Bates et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). For LMMs, p-values were obtained using lmerTest (v3.1.3, Kuznetsova et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), which provides Type III ANOVA tables by default. Beta mixed-effects models were fitted using the glmmTMB package (v1.1.12; Brooks et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Model selection was based on backward elimination of non-significant interaction terms (α\u0026thinsp;=\u0026thinsp;0.05), guided by changes in the Akaike Information Criterion (AIC). In the main text, we report results from the minimal adequate models; full model outputs are provided in the Supplementary Information.\u003c/p\u003e \u003cp\u003eFor clarity, we followed the analytical approach used in recent work (Willcox et al., \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2026\u003c/span\u003e) and collapsed across test sessions, modelling Trial as a single factor with six levels (three trials from each session). The full models\u0026mdash;initially including Test session (session 1 vs. session 2), Trial number (1, 2, or 3), and their interaction to account for between-session differences and within-session learning\u0026mdash;are reported in the Supplementary Information.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eRelationship between MA phenotype and detour task performance in adult females (prediction 1)\u003c/h2\u003e \u003cp\u003eTo test prediction 1, we analysed detour task performance using mixed-effects models. Detour latency was analysed using a linear mixed-effects model with MA phenotype (MA vs noMA) as a fixed effect. Trial number (1\u0026ndash;6) and multi-baseline measure (mean food-approach latency during habituation with an opaque barrier) were included as covariates. Female identity was included as a random intercept to account for repeated measures across trials. Detour latency values were positively skewed; therefore, the response variable was winsorised using a trim value of 0.05 prior to analysis to reduce the influence of extreme values while retaining all observations.\u003c/p\u003e \u003cp\u003ePersistence at the barrier was quantified as the proportion of trial time spent persisting and analysed using a beta mixed-effects model with a logit link function. Because trial duration differed between successful and unsuccessful trials in adult females, a proportional measure was required to ensure comparability across observations. Values of 0 and 1 were adjusted following standard recommendations for beta regression Smithson \u0026amp; Verkuilen, (\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). The same fixed and random effects structure as in the detour latency model was applied.\u003c/p\u003e \u003cp\u003eImprovement in detour latency across trials could be inferred from the MA phenotype \u0026times; Trial interaction term included in the detour latency model described above.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eEffects of early-life exposure to MA on offspring detour task performance (predictions 2a and 2b)\u003c/h2\u003e \u003cp\u003eTo test predictions 2a and 2b, we analysed offspring detour task performance using mixed-effects models that accounted for the cross-fostering design. The MA phenotype of the biological mother and the MA phenotype of the foster mother were included as separate fixed effects, along with their interaction, allowing us to disentangle biological (genetic and prenatal maternal) effects from post-hatching environmental effects. Detour latency was analysed using a linear mixed-effects model with winsorised latency values (trim value\u0026thinsp;=\u0026thinsp;0.05). Trial number and multi-baseline measure were included as covariates. Individual identity nested within nest identity was included as a random intercept to account for repeated measures across trials and non-independence among siblings.\u003c/p\u003e \u003cp\u003eFor offspring, persistence at the barrier was quantified as the total time spent persisting, rather than as a proportion. Because all offspring test trials were of identical duration, absolute persistence times were directly comparable across individuals and trials. Persistence was analysed using a linear mixed-effects model with the same fixed and random effects structure as for detour latency.\u003c/p\u003e \u003cp\u003eImprovement in detour latency across trials could be inferred from the interactions of biological and foster mother MA phenotype with Trial included in the main latency model described above.\u003c/p\u003e \u003cp\u003e \u003cem\u003eAdditional analyses\u003c/em\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003ea) Age-related differences in detour task performance\u003c/h2\u003e \u003cp\u003eBecause inhibitory control develops during ontogeny, we compared juveniles and adults using mixed-effects models, including age class (juvenile vs adult) as a fixed effect. Trial number was included as a covariate. Individual identity was included as a random intercept to account for repeated measures. Detour latency and persistence were analysed as described above. Improvement across trials could be inferred from the Age class \u0026times; Trial interaction included in both models.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003eb) Mother-offspring resemblance in detour task performance\u003c/h2\u003e \u003cp\u003eTo provide exploratory insight into potential intergenerational similarity in inhibitory control, we examined whether offspring detour task performance covaried with the performance of their biological mothers using a mid-offspring regression approach. For each trait, detour latency, mean persistence (proportion of time spent persisting), and improvement across trials, we first calculated a single value per individual by averaging across test sessions. Offspring values were then averaged at the nest level to obtain one mean offspring value per family. These nest-level offspring means were paired with the corresponding maternal values, resulting in one data point per family. This approach avoids pseudo-replication arising from multiple offspring per mother and provides an estimate of intergenerational resemblance at the family level.\u003c/p\u003e \u003cp\u003eWe fitted linear models in which mean offspring performance was used as the response variable and maternal performance as a continuous predictor. For detour latency, both maternal and offspring values were log-transformed prior to analysis. Persistence values were analysed on the original scale. Improvement was calculated as the difference in detour latency between Trial 1 and Trial 6 (Trial 1\u0026thinsp;\u0026minus;\u0026thinsp;Trial 6), such that positive values indicate improved performance across trials. Offspring improvement scores were square-root transformed to meet model assumptions, whereas maternal improvement was entered on the raw scale.\u003c/p\u003e \u003c/div\u003e"},{"header":"RESULTS","content":"\u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003eRelationship between MA phenotype and detour task performance in adult females (prediction 1)\u003c/h2\u003e \u003cp\u003eThe MA phenotype was not associated with detour task performance. Adult females who later expressed MA did not differ from non-aggressive females in their detour latency (\u003cem\u003eMA\u003c/em\u003e: p\u0026thinsp;=\u0026thinsp;0.38; Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea). Across all adult females, detour latency decreased significantly across trials (\u003cem\u003eTrial\u003c/em\u003e: p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), indicating learning over repeated exposures to the task. Adult females who later expressed MA did not spend a different proportion of time persisting at the barrier than females that did not express MA (\u003cem\u003eMA\u003c/em\u003e: p\u0026thinsp;=\u0026thinsp;0.39; Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb). No trial effect was detected for the proportion of time persisting (\u003cem\u003eTrial\u003c/em\u003e: p\u0026thinsp;=\u0026thinsp;0.16). Multi-baseline measure (i.e., mean food-approach latency during habituation with an opaque barrier; see \u003cem\u003eMaterials and methods: Habituation\u003c/em\u003e) did not significantly explain variation in either detour latency or proportion of time persisting (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\u003eOutcome of models testing the association between maternal aggression (MA) phenotype and detour task performance in adult females. Proportion of time persisting was adjusted to accommodate values of 0 and 1. Bold values indicate P\u0026thinsp;\u0026lt;\u0026thinsp;0.05.\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\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cb\u003eDetour Latency\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eEstimate (SE)\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003edf\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 \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIntercept\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.06 (\u0026plusmn;\u0026thinsp;0.15)\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\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMA\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.14 (\u0026plusmn;\u0026thinsp;0.16)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 42.45)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.38\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMulti-baseline measure\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.01 (\u0026plusmn;\u0026thinsp;0.01)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 46.42)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.69\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrial\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.20 (\u0026plusmn;\u0026thinsp;0.02)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e204.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 204.81)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\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\u003e\u003cb\u003eEstimate (SE)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003e\u003cb\u003eZ value\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eProportion of time persisting\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\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\u003e\u003cem\u003eIntercept\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.28 (\u0026plusmn;\u0026thinsp;0.18)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003e1.53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.12\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMA\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.14 (\u0026plusmn;\u0026thinsp;0.17)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003e-0.85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.39\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMulti-baseline measure\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.01 (\u0026plusmn;\u0026thinsp;0.01)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003e0.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.77\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrial\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.05 (\u0026plusmn;\u0026thinsp;0.03)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003e-1.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.16\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003eEffects of early-life exposure to MA on offspring detour task performance (prediction 2a and 2b)\u003c/h2\u003e \u003cp\u003eThere was no evidence that early-life exposure to MA was associated with offspring detour task performance. Neither post-hatching exposure to MA (\u003cem\u003efoster mother phenotype\u003c/em\u003e: p\u0026thinsp;=\u0026thinsp;0.59) nor the MA phenotype of the biological mother (\u003cem\u003ebiological mother phenotype\u003c/em\u003e: p\u0026thinsp;=\u0026thinsp;0.13; Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea) significantly explained variation in offspring detour latency. Similarly, total time spent persisting at the barrier did not differ according to foster (\u003cem\u003efoster mother phenotype\u003c/em\u003e: p\u0026thinsp;=\u0026thinsp;0.73) or biological (\u003cem\u003ebiological mother phenotype\u003c/em\u003e: p\u0026thinsp;=\u0026thinsp;0.89) mother MA phenotype (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb).\u003c/p\u003e \u003cp\u003eAcross offspring, detour latency and total time persisting both decreased significantly across trials (\u003cem\u003eTrial\u003c/em\u003e: p\u0026thinsp;\u0026lt;\u0026thinsp;0.001 in both cases), indicating learning over repeated exposures to the task. The multi-baseline measure was a significant covariate for detour latency (p\u0026thinsp;=\u0026thinsp;0.04; Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e): individuals with longer food-approach latencies during habituation also showing longer detour latencies during testing. It did not significantly predict total time persisting (see Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\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\u003eOutcome of mixed-effects models testing the relationship between maternal aggression (MA) exposure and detour task performance in offspring. Detour latency and total time persisting were analysed using linear mixed-effects models. Detour latency was winsorised (trim\u0026thinsp;=\u0026thinsp;0.05). Groups were defined by the MA phenotype of the biological and foster mothers. Bold values indicate P\u0026thinsp;\u0026lt;\u0026thinsp;0.05.\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\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cb\u003eDetour latency\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eEstimate (SE)\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003edf\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 \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIntercept\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.96 (\u0026plusmn;\u0026thinsp;0.12)\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\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMulti-baseline measure\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.01 (\u0026plusmn;\u0026thinsp;0.01)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 105.72)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e0.04\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eBiological mother phenotype\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.22 (\u0026plusmn;\u0026thinsp;0.14)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 108.59)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.13\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eFoster mother phenotype\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.08 (\u0026plusmn;\u0026thinsp;0.15)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 111.73)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.59\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrial\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.27 (\u0026plusmn;\u0026thinsp;0.01)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e306.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 566.51)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\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\u003e\u003cb\u003eEstimate (SE)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003edf\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal time persisting\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIntercept\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e8.80 (\u0026plusmn;\u0026thinsp;0.85)\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\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMulti-baseline measure\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.01 (\u0026plusmn;\u0026thinsp;0.01)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 94.81)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.27\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eBiological mother phenotype\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.11 (\u0026plusmn;\u0026thinsp;0.93)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 89.30)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.89\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eFoster mother phenotype\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.33 (\u0026plusmn;\u0026thinsp;0.98)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 108.05)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.73\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrial\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-1.64 (\u0026plusmn;\u0026thinsp;0.17)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e91.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 571.10)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec21\" class=\"Section2\"\u003e \u003ch2\u003eAge-related differences in detour task performance\u003c/h2\u003e \u003cp\u003eDetour task performance differed markedly between juveniles and adults. For detour latency, there was a strong main effect of age class (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea), with juveniles showing longer detour latencies than adults. Latency decreased significantly across trials (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), indicating overall improvement in task performance. Importantly, the interaction between age class and trial number was significant (p\u0026thinsp;=\u0026thinsp;0.04), showing that the rate of improvement differed between age classes, with juveniles exhibiting a steeper decline in latency across trials than adults.\u003c/p\u003e \u003cp\u003eA similar pattern emerged for persistence at the barrier. There was a significant effect of age class (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Table\u0026nbsp;4; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb), with juveniles spending less time persisting at the barrier than adults. Persistence decreased across trials (p\u0026thinsp;=\u0026thinsp;0.02), consistent with improved task performance over time. The interaction between age class and trial was also significant (p\u0026thinsp;=\u0026thinsp;0.02), indicating that juveniles showed a steeper reduction in persistence across trials, while adult persistence remained comparatively stable, consistent with the absence of a significant trial effect for adult persistence observed in the prediction 1 analyses (\u003cem\u003eTrial\u003c/em\u003e: p\u0026thinsp;=\u0026thinsp;0.16; see Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\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\u003eOutcome of models testing the association between age class and detour task performance. Detour latency and proportion of time persisting were analysed using linear mixed-effects models. Detour latency was log-transformed and winsorised (trim\u0026thinsp;=\u0026thinsp;0.05). Proportion of time persisting at the barrier was square-root transformed. Bold values indicate \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05.\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\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cb\u003eDetour latency\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eEstimate (SE)\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003edf\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 \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIntercept\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.13 (\u003cem\u003e\u0026plusmn;\u003c/em\u003e\u0026thinsp;0.09)\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\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAge class\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.82 (\u003cem\u003e\u0026plusmn;\u003c/em\u003e\u0026thinsp;0.12)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e41.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 511.22)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrial\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.22 (\u003cem\u003e\u0026plusmn;\u003c/em\u003e\u0026thinsp;0.01)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e134.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 985.90)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAge class: Trial\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.04 (\u003cem\u003e\u0026plusmn;\u003c/em\u003e\u0026thinsp;0.02)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 963.89)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e0.04\u003c/b\u003e\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\u003e\u003cb\u003eEstimate (SE)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eF\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003edf\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eProportion of time persisting\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIntercept\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.16 (\u003cem\u003e\u0026plusmn;\u003c/em\u003e\u0026thinsp;0.06)\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\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAge class\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.61 (\u003cem\u003e\u0026plusmn;\u003c/em\u003e\u0026thinsp;0.11)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e31.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 170.98)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrial\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.04 (\u003cem\u003e\u0026plusmn;\u003c/em\u003e\u0026thinsp;0.01)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4.89\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 268.32)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e0.02\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAge class: Trial\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.09 (\u003cem\u003e\u0026plusmn;\u003c/em\u003e\u0026thinsp;0.04)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1, 282.24)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e0.02\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec22\" class=\"Section2\"\u003e \u003ch2\u003eMother-offspring resemblance in detour task performance\u003c/h2\u003e \u003cp\u003eWe found no evidence for mother-offspring resemblance in inhibitory control. Mean offspring detour latency was not associated with maternal detour latency (β = \u0026minus;0.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14 SE, \u003cem\u003eF\u003c/em\u003e₁, ₂₉ = 0.75, p\u0026thinsp;=\u0026thinsp;0.39; Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea), indicating that offspring from mothers with slower or faster detour performance did not differ in their own latency to solve the task.\u003c/p\u003e \u003cp\u003eSimilarly, offspring persistence at the barrier was not related to maternal persistence (β = \u0026minus;0.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08 SE, \u003cem\u003eF\u003c/em\u003e₁, ₂₉ = 0.41, p\u0026thinsp;=\u0026thinsp;0.53; Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eb), providing no evidence for intergenerational similarity in this behavioural component of the task.\u003c/p\u003e \u003cp\u003eConsistent with these findings, improvement in detour performance across trials was not associated with maternal improvement (β = -0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07, \u003cem\u003eF\u003c/em\u003e₁, \u003csub\u003e22.63\u003c/sub\u003e = 1.30, p\u0026thinsp;=\u0026thinsp;0.26; Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec), indicating that offspring did not resemble their mothers in their rate of performance change over repeated trials.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"DISCUSSION","content":"\u003cp\u003eThis study examined whether maternal aggression (MA) is associated with individual differences in inhibitory control in adult canary females and whether early-life exposure to MA contributes to variation in offspring inhibitory control. Contrary to our predictions, we found no evidence that MA phenotype in adult females was associated with detour task performance, indicating that aggressive and non-aggressive mothers did not differ in their ability to inhibit prepotent motor responses. Despite representing a harsh early-life social environment, post-hatching exposure to MA did not influence offspring inhibitory control. This absence of environmental effects is notable given that executive functions are often considered sensitive to early developmental conditions. Detour performance varied with age: juveniles showed longer detour latencies than adults and improved more steeply across trials, suggesting that behavioural performance in this task continues to develop after hatching. However, adults spent more time persisting at the barrier than juveniles. Finally, we found no evidence for mother-offspring resemblance in detour task performance.\u003c/p\u003e \u003cdiv id=\"Sec24\" class=\"Section2\"\u003e \u003ch2\u003eMaternal aggression is not associated with impaired inhibitory control in adult females\u003c/h2\u003e \u003cp\u003eThe absence of an association between MA and detour task performance in adult females suggests that MA in canaries is unlikely to reflect reduced response inhibition or heightened impulsivity, at least as captured by this task. While some studies have reported links between aggression and inhibitory control in birds, most notably in Australian magpies (\u003cem\u003eGhymnorhina tibicen\u003c/em\u003e), where aggressive or risk-prone individuals performed more poorly in tasks requiring motor control or behavioural flexibility (Speechley et al., \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; but see also, Vernouillet et al., \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), our findings align with a growing body of work that fails to replicate such associations. For example, van Iersel et al. (\u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) found no relationship between territorial aggression and detour task performance in free-living female blue tits (\u003cem\u003eCyanistes caeruleus\u003c/em\u003e), and Willcox et al., (\u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2026\u003c/span\u003e) similarly reported no association between aggression and response inhibition in chickens (\u003cem\u003eGallus gallus domesticus\u003c/em\u003e). Along with our study, these recent findings challenge the assumption of a generalised cognitive-behavioural syndrome linking aggression to impaired inhibition. However, detour tasks typically involve not only response inhibition, but also problem-solving, spatial learning, and exploration, meaning that environment-dependent effects on inhibitory control specifically may be diluted at the task level (Kabadayi et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; van Horik et al., \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Thus, the measures we obtained are composite, and this may limit their sensitivity to specific regulatory processes. Moreover, performance often shows low consistency across different inhibition tasks, suggesting that task design strongly influences which inhibitory processes are captured (Brucks et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Troisi et al., \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2026a\u003c/span\u003e, \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2026b\u003c/span\u003e; van Horik et al., \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; V\u0026ouml;lter et al., \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). This should be borne in mind when interpreting the present study. That said, task-level limitations alone cannot fully account for the absence of an association between MA and detour performance, since they would equally apply to studies that do detect such links between aggression and inhibitory control. A more fundamental explanation may therefore lie in the nature of MA itself.\u003c/p\u003e \u003cp\u003eMA in canaries may not represent reactive or impulsive aggression, but rather a context-dependent component of a parental investment strategy. From this perspective, MA may represent an escalation of selective investment decisions rather than deficits in behavioural regulation. Comparable forms of parental aggression or brood reduction have been documented in birds such as coots (\u003cem\u003eFulica atra)\u003c/em\u003e, storks (\u003cem\u003eCiconia ciconia\u003c/em\u003e), and pelicans (\u003cem\u003ePelecanus erythrorhynchos\u003c/em\u003e), where aggression towards offspring facilitates brood reduction under limited resources (see Cash \u0026amp; Evans, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Shizuka \u0026amp; Lyon, \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Tortosa \u0026amp; Redondo, \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Zieliński, \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), as well as in mammals, where maternal rejection or aggression biases investment towards higher-quality offspring (e.g., Maestripieri, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Mock \u0026amp; Parker, \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). If MA serves as a strategic adjustment rather than an impulsive response, it would not be expected to co-vary with individual differences in inhibitory control measured in a non-social, non-breeding context.\u003c/p\u003e \u003cp\u003eInterestingly, MA females did not score higher than noMA females on a social dominance test conducted outside the breeding context (Garcia-Co et al., in prep.), suggesting that MA does not reflect a broadly aggressive phenotype. Rather, aggressiveness in the breeding context appears to be decoupled from social behaviour in other contexts, consistent with the view that MA is a state-dependent, context-specific trait (Dingemanse et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; R\u0026eacute;ale et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Aggression is a multi-faceted behaviour, regulated by distinct neural and endocrine mechanisms depending on its functional context (e.g., territorial, social, or parental), and individuals often express aggression selectively rather than consistently across situations (Dingemanse et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Maney \u0026amp; Goodson, \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Nelson \u0026amp; Trainor, \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Yanowitch \u0026amp; Coccaro, \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). If MA is not part of a broader, cross-context aggressive phenotype, then it is unlikely to be associated with individual differences in inhibitory control measured in a non-social, non-reproductive setting. Together, these considerations suggest that the absence of an association between MA and detour performance reflects the context-specificity of MA expression, rather than a genuine absence of any link between inhibitory control and aggressive behaviour in this species.\u003c/p\u003e \u003cdiv id=\"Sec25\" class=\"Section3\"\u003e \u003ch2\u003eEarly-life exposure to maternal aggression does not alter offspring inhibitory control\u003c/h2\u003e \u003cp\u003eWe found no evidence that offspring raised by aggressive mothers differed in detour task performance from offspring associated with non-aggressive mothers. This absence of post-hatching environmental effects is notable given that MA constitutes an early-life social stressor and has been linked to long-term behavioural differences in this system, including heightened aggressiveness and reduced neophobia (Garcia-Co et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2025\u003c/span\u003e, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2026\u003c/span\u003e). Furthermore, a substantial body of work shows that early-life conditions can shape inhibitory control and related executive functions across taxa (Brett et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Dydenkova et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Hedges \u0026amp; Woon, \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Rowell et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In birds, for example, experimental manipulations of early life environmental predictability, enrichment, and social conditions have been shown to influence inhibitory control and persistence in detour or cylinder tasks (Ashton et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; van Horik et al., \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Vernouillet et al., \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Willcox et al., \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2025\u003c/span\u003ea), although such effects are not consistently detected (Willcox et al., \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2026\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeveral considerations may explain this discrepancy. First, as noted above, detour performance is a composite measure while early-life MA may selectively affect inhibitory processes without this being detectable at the level of overall detour performance. Indeed, in a previous study examining responses to an escape test (Garcia-Co et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2026\u003c/span\u003e), offspring born to biological mothers that later exerted MA spent more time pecking at a closed barrier, a measure of persistence, than nestlings born to non-aggressive biological mothers. This suggests that early-life MA does shape some aspects of behavioural regulation, but that these effects may be domain-specific and not generalise across task contexts. Effects may also be more apparent in socially embedded tasks where the inhibitory demands most relevant to an adverse early social environment are more directly engaged. Consistent with this, Willcox et al., (\u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2026\u003c/span\u003e) found that early-life social instability in chickens affected aggression in a social context but not response inhibition in a non-social context.\u003c/p\u003e \u003cp\u003eSecond, we observed strong age-related differences in detour task performance, with juveniles showing longer detour latencies than adults but improving more steeply over trials, consistent with a gradual maturation of executive functions. The steeper improvement in juveniles likely reflects greater room for improvement at this developmental stage rather than higher learning capacity per se, as individuals starting from a lower baseline have more potential for short-term gain. Comparable ontogenetic trajectories are well documented across birds and mammals, with inhibitory control improving as neural circuits mature and individuals accumulate experience (Diamond, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Diamond \u0026amp; Gilbert, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1989\u003c/span\u003e; Kabadayi et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Canaries are an altricial species, meaning that neural development continues well after hatching (Iwaniuk \u0026amp; Nelson, \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). Our juveniles were tested shortly after independence, at a stage when inhibitory circuits may not have been fully established at the juvenile stage (Diamond, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Kabadayi et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). If the neural systems underlying detour performance are still maturing at the time of testing, they may also be less responsive to the influence of earlier social experience, potentially masking effects of early-life MA that might only emerge later in development. Indeed, in some species the effects of early social environment on cognitive performance have been detected only later in ontogeny the effects of early social environment on cognitive performance emerged only later in ontogeny (Ashton et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; but see van Horik et al., \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Vernouillet et al., \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Willcox, et al., \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Whether MA-exposed offspring would show differences in inhibitory control once fully mature remains an open question.\u003c/p\u003e \u003cp\u003eInterestingly, the pattern for persistence was the reverse: adults spent more time persisting at the barrier than juveniles throughout testing. This could reflect greater goal-directedness or motivation in adults, as mature individuals may invest more effort in a direct approach before switching strategies, whereas juveniles might disengage more rapidly after an unsuccessful attempt. Notably, despite their higher persistence, adults still showed generally fast detour latencies overall, suggesting that the time spent near the barrier did not substantially delay task success. This may indicate that repeated barrier approaches carry a low behavioural cost for adults in this context (e.g., Kabadayi et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; van Horik et al., \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), and that adult persistence does not necessarily reflect an inhibitory deficit but rather a different pattern of task exploration. Furthermore, juveniles showed a steeper reduction in persistence across trials than adults, mirroring the steeper improvement in detour latency. Whether this reflects greater neural plasticity or simply lower initial motor fixation on the barrier remains to be determined. Regardless of the underlying mechanism, these results suggest that while inhibitory control improves with age, other motivational factors, such as persistence, may follow a distinct developmental trajectory.\u003c/p\u003e \u003cp\u003eWe found no evidence for mother-offspring resemblance in detour task performance, suggesting limited genetic or prenatal maternal effects in this measure of inhibitory control. This is consistent with low heritability estimates reported for inhibitory control and related cognitive traits in birds (Gnanadesikan et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Langley et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Together with the lack of early environmental effects, the absence of genetic effects supports the view that inhibitory control measures are among others shaped by an individual's experience and current motivational state (Bateson et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Dunn et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019a\u003c/span\u003e). Limited resemblance may additionally reflect the high plasticity of inhibitory control as a trait. Measures that are strongly responsive to individual experience and current context are inherently more difficult to establish as heritable from a limited number of observations per individual (Brucks et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; McCallum \u0026amp; Shaw, \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Troisi et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e"},{"header":"Conclusions","content":"\u003cp\u003eOverall, our results suggest that MA in canaries is not associated with a general impairment in inhibitory control and does not appear to be transmitted across generations through variation in this cognitive trait. The absence of an association between MA and detour performance, together with the lack of cross-context consistency in aggressive behaviour, suggests that MA is a state-dependent, context-specific maternal behaviour rather than the expression of a stable inhibitory deficit. Evidence from a previous study in this population (Garcia-Co et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2026\u003c/span\u003e) nevertheless suggests that early-life MA does selectively shape some aspects of behavioural regulation, pointing to domain-specific effects.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eDATA AVAILABILITY\u003c/p\u003e\n\u003cp\u003eThe data used in this study is archived in Mendeley Data in the following link: https://data.mendeley.com/datasets/cy8fhzsy56/1. The videos analysed are available from the corresponding author (Clara Garcia-Co) upon request.\u003c/p\u003e\n\u003cp\u003eAUTHOR CONTRIBUTIONS\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eClara Garcia-Co:\u003c/strong\u003e Data curation; Formal analysis; Investigation; Methodology; Validation; Visualization; Writing \u0026ndash; original draft \u0026amp; editing. \u0026nbsp;\u003cstrong\u003eAnneleen Dewulf:\u003c/strong\u003e Data curation; Formal analysis; Investigation; Methodology; Validation; Writing \u0026ndash; review \u0026amp; editing. \u003cstrong\u003eJudith Morales:\u003c/strong\u003e Funding acquisition; Supervision; Validation; Writing \u0026ndash; review \u0026amp; editing. \u003cstrong\u003eFrederick Verbruggen:\u003c/strong\u003e Funding acquisition; Supervision; Validation; Writing \u0026ndash; review \u0026amp; editing. \u0026nbsp;\u003cstrong\u003eWendt M\u0026uuml;ller:\u003c/strong\u003e Conceptualization; Funding acquisition; Investigation (supporting); Methodology (supporting); Resources; Supervision; Validation; Writing \u0026ndash; review \u0026amp; editing.\u003c/p\u003e\n\u003cp\u003eCONFLICT OF INTEREST STATEMENT\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing or financial interests.\u003c/p\u003e\n\u003cp\u003eACKNOWLEDGEMENTS\u003c/p\u003e\n\u003cp\u003eWe thank Peter Scheys for his assistance in taking care of the birds.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFUNDING STATEMENT\u003c/p\u003e\n\u003cp\u003eThis work was supported by the FWO Flanders (CG: 1173825N and AD: 11F0823N). FV was supported by an ERC Consolidator grant (European Union\u0026rsquo;s Horizon 2020 research and innovation programme, grant agreement No 769595) and Methusalem Project 01M00221 (Ghent University). JM was supported by PID2022-139166NB-I00 funded by MCIN/AEI/https://doi.org/10.13039/501100011033 and \u0026ldquo;ERDF A way of making Europe\u0026rdquo;.\u003c/p\u003e\n\u003cp\u003eSTATEMENT OF ANIMAL ETHICS\u003c/p\u003e\n\u003cp\u003eThe experiments described in this study were carried out with the approval of the Ethical Committee for Animal Experimentation of the University of Antwerp (ECD 2022-87) and by the Ethical Committee for Animal Experimentation, Faculty of Sciences, Flemish Institute for Biotechnology (VIB) of Ghent University (EC2022-091). We complied with Belgian and Flemish animal welfare legislation, adhered to the ASAB/ABS guidelines for the use of animals in behavioural research and teaching, and ARRIVE guidelines. A constant effort was made to keep the birds in semi-natural conditions and in good health. The invasiveness of all observational procedures described was minimized, and handling time was kept under two minutes per bird. Birds were handled exclusively by trained personnel (AD, CG, WM: FELASA C; caretakers: FELASA B).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eBody mass and behaviour were monitored throughout the whole experiment, as they reflect general well-being (see Paul-Murphy \u0026amp; Hawkins, 2015). Nestlings were temporarily marked with non-toxic coloured markers until they reached approximately 7 g, when they were fitted with standard metal leg rings; these methods have negligible effects in passerines (Eldegard et al., 2024). For short-term visual identification in group tests, a very small (\u0026lt; 2 mm) dot of correction fluid was applied to crown feathers only. This method is consistent with field marking guidance and produces no observable irritation (Department of Biodiversity, Conservation and Attractions, 2020; NORECOPA, 2022). For food-motivated tasks, birds were food-restricted overnight (for a maximum of 18 hours) to standardise motivation across individuals; this mild deprivation is within natural foraging intervals (see e.g., Cauchoix et al., 2017; Dunn et al., 2019a; Vergauwen et al., 2014). At the end of the experiment, all the canaries were kept in the lab-bred population at the University of Antwerp as they are part of a transgenerational transmission experiment and were allowed to breed in the following years.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAnda RF, Felitti VJ, Bremner JD, Walker JD, Whitfield C, Perry BD, Dube SR, Giles WH (2006) The enduring effects of abuse and related adverse experiences in childhood. 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Anim Cogn 8(2):122\u0026ndash;128. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10071-004-0243-x\u003c/span\u003e\u003cspan address=\"10.1007/s10071-004-0243-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\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":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"animal-cognition","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"anco","sideBox":"Learn more about [Animal Cognition](http://link.springer.com/journal/10071)","snPcode":"10071","submissionUrl":"https://submission.nature.com/new-submission/10071/3","title":"Animal Cognition","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"inhibitory control, response inhibition, maternal aggression, detour task, transgenerational transmission, ontogeny","lastPublishedDoi":"10.21203/rs.3.rs-9402003/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9402003/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eMaternal aggression (MA) towards offspring represents an early-life social stressor and has been linked to long-term behavioural differences. Yet, the cognitive mechanisms underlying its expression and potential intergenerational transmission remain unclear. We tested whether MA in domestic canaries (\u003cem\u003eSerinus canaria\u003c/em\u003e) is associated with individual differences in inhibitory control (i.e., the ability to suppress impulsive or inappropriate actions) in adult females and whether early-life exposure to MA subsequently predicts variation in offspring inhibitory control. Inhibitory control was assessed using a detour task, in which individuals must suppress a direct approach to a visible reward in favour of an indirect solution. A cross-fostering design was used to disentangle post-hatching environmental effects from prenatal or genetic influences on offspring performance. We found no evidence that aggressive and non-aggressive mothers differed in inhibitory control. Likewise, offspring inhibitory control was not influenced by post-hatching exposure to MA nor by the MA phenotype of the biological mother, and we detected no mother\u0026ndash;offspring resemblance in inhibitory control. However, strong ontogenetic effects were observed: juveniles showed longer detour latencies than adults but improved more steeply across trials, consistent with a post-hatching maturation of inhibitory control. Together, these results suggest that MA in this system is unlikely to reflect deficits in inhibitory control or to be transmitted across generations via this cognitive pathway. More broadly, our findings highlight the task- and context-specific nature of inhibitory control measures.\u003c/p\u003e","manuscriptTitle":"No evidence for a link between maternal aggression and inhibitory control across generations in domestic canaries","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-29 11:03:54","doi":"10.21203/rs.3.rs-9402003/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"37597059126588920201955731450076236718","date":"2026-05-13T02:55:56+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-04-20T20:05:17+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-04-15T18:16:03+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-04-15T08:09:52+00:00","index":"","fulltext":""},{"type":"submitted","content":"Animal Cognition","date":"2026-04-13T09:26:20+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"animal-cognition","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"anco","sideBox":"Learn more about [Animal Cognition](http://link.springer.com/journal/10071)","snPcode":"10071","submissionUrl":"https://submission.nature.com/new-submission/10071/3","title":"Animal Cognition","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"b1728253-2b8e-40b0-90b8-ea1a12c32114","owner":[],"postedDate":"April 29th, 2026","published":true,"recentEditorialEvents":[{"type":"reviewerAgreed","content":"37597059126588920201955731450076236718","date":"2026-05-13T02:55:56+00:00","index":24,"fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-04-29T11:03:54+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-29 11:03:54","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9402003","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9402003","identity":"rs-9402003","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}