Elucidating the role of neurotransmitters in the behavioural plasticity of Camponotus compressus (Hymenoptera: Formicidae)

Elucidating the role of neurotransmitters in the behavioural plasticity of Camponotus compressus (Hymenoptera: Formicidae) | 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 Elucidating the role of neurotransmitters in the behavioural plasticity of Camponotus compressus (Hymenoptera: Formicidae) Swasti Saxena, Ankita Salunke, Nishi Pandya This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8751427/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Ants express a remarkable behavioural plasticity ranging far beyond rigid stimulus-response relationships. They are omnivorous and each caste exhibit a variety of behaviour including nesting (foraging, food storage, protection from predators, feeding, nest structure and brood care). The behaviour investment of workers is shaped by presence or absence of queen within the colony; which may affect the behavioural flexibility and developmental neuroplasticity in workers. However, there is a lacuna in the studies conducted in the nesting behaviour and the associated neurophysiology of such ants. So, the present study was aimed to determine the role of neurotransmitters (Dopamine and Serotonin) underlining neuroplasticity in the nesting behaviour−foraging, feeding, searching, immobility, communication, defence, and nest structure of Camponotus compressus ant, under laboratory conditions. Our results demonstrated an increase in the respective behavioural activities, as well as in the length and depth of the nest, in the presence of the queen, with a subsequent decrease on the 14th day following her removal. Additionally, neurotransmitter levels (DA and 5-HT) were higher in workers (media and minor) ants on the 7th day, followed by a significant ( p < 0.05 ) decline after 14th day suggesting a reduced nesting activity, which is further correlated with the decrease in nest morphometry, in comparison to control. The current study unveils the neurophysiological mechanism involved in achieving neuroplasticity in the C. compressus . Further studies on specific gene expressions and histochemistry of brain will provide a broader view on the altered nesting behaviour in worker ants and the developed plastic queen. Camponotus compressus Nesting Behaviour Neuroplasticity Dopamine Serotonin Figures Figure 1 Figure 2 Figure 3 Introduction Animal personality and behavioural syndromes, conflated in earlier literature (MacKay and Haskell 2015 ; Sánchez-Tójar et al. 2022 ), represents distinct conceptual frameworks: animal personality describes individual consistency in specific traits across contexts including aggression stability across ecological fluctuations (Bell and Sih 2007 ), while behavioural syndromes like boldness-aggressiveness and associations address traits correlations within individuals (Sih et al. 2004a , b ). Some studies have demonstrated the absence of significant correlations between behavioural traits (Carvalho et al. 2013 ; Roth et al. 2021 , 2022 ). Insects, frequently regarded as less adaptable than vertebrates (Chittka and Niven 2009 ; Wystrach and Graham 2012 ), typically exhibit compensatory abilities for impairments as immediate responses, attributed to the inherent natural redundancy and robustness of their systems, rather than as outcomes of lifelong neural plasticity (Wystrach and Graham 2012 ; Stone et al. 2022 ). Insects like ants, cockroaches, stick insects and other insects, for instance, show a robust and spontaneous neural response governing leg coordination (Steck et al. 2009 ; Schwarz et al. 2024 ). Insect colonies are formed of a number of individuals each representing a distinct labour such as reproductive division of tasks between queen, king (in termites), largely sterile workers as well as communication between and within the caste for the efficient functioning of a colony, essential for ecological success (Hӧlldobler and Wilson 1990). Workers respond to the changing signals in a way that are beneficial for the entire colony (Hӧlldobler and Wilson 2009). Ants, including largely of castes—queen, drone (male), and worker ants (Clark and Holbrook 2009 ) exhibit a distinctive nesting pattern characterised by social interaction in nest construction. Communication within the colony adheres to a hierarchical framework, wherein minor ants predominantly engage with medium ants, who subsequently convey information to the queen (Wollman et al. 2011). Consequently, they construct a nest featuring intricate, network-like structures composed of chambers interconnected by tunnels that serve as nodes and connecting links, respectively (Tschinkel 2004 ; Buhl et al. 2004 ). Improved communication among chambers can accelerate resource transfer inside the nest (Cook et al. 2014 ). Members of a colony are divided into castes with distinct characteristics, such as behaviour, metabolism, lifespan, and reproduction. Only a single queen is accountable for reproduction, possessing ovarioles with oocytes at various stages of differentiation (Khila and Abouheif 2008 ; Lee et al. 2017 ; Penick et al. 2021 ). Conversely, workers are non-reproductive, possessing undeveloped ovaries which constitute that exhibit distinct characteristics from the queen, engaging in incessant movement, brood care, colony sanitation, and food foraging, which are acquired during the embryonic and larval developmental stages (Khila and Abouheif 2010 : Schwander et al. 2010 ; Trible and Kronauer 2021). Queen signalling is crucial for ant colonies as it signals the queen's presence and reproductive status, prompting workers to forgo their own reproductive efforts and focus on assisting with brood care (Keller and Nonacs 1993 ; Leonhardt et al. 2016 ). Moreover, worker ants signal and encourage others to follow them to the food source through tandem running (Witte et al. 2007 ; Holldobler and Wilson 2009; Simpson and Raubenheimer 2012 ; Csata and Dussutour 2019 ). Female workers typically do not reproduce; instead, they are responsible for colony maintenance and brood care. However, despite this clear division of roles, workers can become reproductively active under certain conditions. The presence of a queen strongly suppresses this reproductive flexibility. In queenless species, colony reproduction is managed through a well-organized system where reproductive and non-reproductive tasks are distinctly divided among workers. The reproductive ability in a queenless colony lies to some individuals of the colony, for example, in Neoponera apicalis , the workers establish a hierarchy based on the behaviours such as antennal dueling and biting. One of the dominant higher worker which exhibits a higher frequency of this behaviour, lays most of the eggs which result into male progenies. Also, in Harpegnathos saltator , the colony is controlled by one queen or a bunch of gamergates which is a mated worker, with a sexually reproducing spermatheca (Hölldobler and Wilson 1990 ; Peeters and Hölldobler 1995 ; Monnin and Peeters 1999 ; Aldana et al. 2024 ). Interpreting the concept of regulation in social insects poses a distinct challenge, especially in comprehending the neurological mechanisms that link behavioural conditioning in the brain to alterations in reproductive physiology. Biogenic amines act as neurotransmitters, neuromodulators, and neurohormones in the brain, playing a vital role in the regulation of behaviour (Roeder 2005 ; Scheiner et al. 2006 ; Pandya et al. 2022 ). In insects, the competition for reproductive hierarchy occurs among the colony’s females with behavioural consequences frequently regulated by neurochemical pathways. Moreover, the concentration of biogenic amines in brain tissue can profoundly influence individual worker behaviour by modulating their responsiveness to environmental inputs, including foraging (Friedman et al. 2018 ). Serotonin and dopamine, as biogenic amines, are recognized for their role in regulating dominance-related behaviours, particularly aggression among colony caste members (Miczek et al. 2002 ; Kravitz and Huber 2003 ; Nelson 2006 ). Octopamine and serotonin are associated with aggression in crickets (Adamo et al. 1995 ; Murakami and Itoh 2001 ; Dyakonova et al. 2002 ; Barbero et al. 2023 ); dopamine has been linked to social dominance in both vertebrates and solitary insects (Baier et al. 2002 ; Miczek et al. 2002 ; Stevenson et al. 2005 ). Biogenic amines regulate key aspects of ant behaviour, with serotonin and dopamine modulating aggression, defense, and arousal, while octopamine and tyramine influence social interactions and aggression control. Additionally, shifts in amino acid ratios with age and task specialization underscore their role in coordinating both individual and collective colony functions (Seid and Traniello 2005 ; Smith et al. 2013; Szczuka 2013; Koyama et al. 2015 ; Aonuma 2020 ). Biogenic amines modulates aggression in Formica polyctena ants, wherein exogenous administration of serotonin, dopamine, octopamine, and tyramine was assessed during dual aggression tests involving nest mates, alien conspecifics, allospecific ants ( Formica fusca ), and prey (house cricket nymphs, Acheta domesticus ) (Vander Meer et al. 2008; Ohkawara and Aonuma 2016 ; Shimoji et al. 2017 ). Serotonin gradually enhances physical aggression, encompassing biting and the secretion of formic acid whereas dopamine increases the mandible threats and biting toward both F. fusca and cricket nymphs (Szczuka et al. 2013 ). On the other hand, octopamine exhibited no substantial impact on aggressive behaviours. Such variations in reproductive status and behaviour among nestmates in ant and bee species are further associated with tissue-specific physiological and transcriptomic differences (Jandt and Gordon 2016 ; Jeanne 2016 ; Toth and Dolezal 2017 ; Chandra et al. 2018 ). In particular, biogenic amine and neurohormonal signaling in the brains of workers are crucial for regulating foraging behaviour in both social and solitary insects. Alterations in brain biogenic amine concentrations can influence individual workers’ responses to specific stimuli, including foraging cues (Muscedere et al. 2012; Kamhi and Traniello 2013 ; Bubak et al. 2016 ; Scheiner et al. 2017 ). Natural differences in nestmates' sensitivity to stimuli can be beneficial for colony function, such as enabling flexible task allocation. Ants are considered a highly suitable model organism for exploring the neurobiological studies of foraging behaviour of group animals to account the highly organized structures built due to the collective efforts of the colony (Li et al. 2014 ). Studies suggest that elimination of queen may cause imbalance in the neurotransmitter levels and behavioural activities which might result into neural plasticity. However, there is a lacuna in the studies conducted on behavioural plasticity of commonly found ants like the C. compressus . Therefore, the present study was aimed to elucidate the role of neurotransmitters in plasticity of C. compressus ants. We assessed the behavioural plasticity in worker ants by evaluating the nesting behaviour, levels of neurotransmitters and whether a worker ant achieves the plasticity or not. Materials and Methods Collection, rearing and acclimatisation Camponotus compressus were collected from the grounds of the Pavilion of Maharaja Sayajirao University of Baroda (22.320153 o N, 73.185734 o E and 22.31992 o N, 73.185851 o E) of Vadodara city, India. Collection of C. compressus was carried out during the noon in the months of December 2024 to May 2025, using handpicking method from the ground and bark of the tree. The collected ant colonies were brought to laboratory for identification and rearing. Morphological identification was done up to the species level with the help of standard taxonomic keys and by comparing the specimens in department repository (Sheikh et al. 2017 ). The ants−30 minor, 19 media, and 1 queen were reared in nesting and foraging boxes (3 sets, sized-15.0 ⋅ 8.5 ⋅ 20 cm) under laboratory conditions (26 ° C temperature, 75% humidity and 12h/12h night and day cycles), using the method of Soares et al. ( 2008 ) and Heinze 2020 (Fig. 1). They were fed with supplements like sugar solution (4mL of 40% solution) for the carbohydrate content while freshly dead or alive but crushed insects for the protein requirement of the ants (Szabó et al. 2025 ). The dead ants were removed from the setup to eliminate any negative effects on the rearing and maintenance. The soil required for the rearing was collected from the same sampling sites at the time of ant collection in order to provide a suitable environment (Czechowski and Pisarski 1992 ). Soil Analysis The physical and chemical parameters of the collected soil were checked to analyse the type of soil and its quality in relation to C. compressus. Soil texture identification was done by texture analysis and further, the soil porosity, water holding capacity and pH were measured using the method given by Saxena ( 2001 ). Behavioural studies Experimental setup After acclimatisation period of the ants in the control and experimental setup (30 minor, 19 media and 1 queen for each group), the behavioural activities of C. compressus were observed and recorded on 7th ,14th, 21st, 28th, and 35th day. In the experimental setup, the queen ant was removed on the 7th day and the worker ants were observed for their behaviour and neural regulation in absence of queen. Food was provided at regular intervals to ensure the normal functioning and behaviour of C. compressus . Behaviour Several behavioural patterns such as tunneling, feeding, foraging, searching, immobility, communication, and defence were examined across the experimental and control group. For each behaviour, readings were taken after every four days, with each session lasting five hours comprising of one-hour intervals between every observation. The ant colonies were observed regularly to analyze acclimatisation to the laboratory conditions. The behavioural reservoire of C. compressus under laboratory conditions was assessed through multiple parameters. Tunneling behaviour was recorded by measuring the number, length, and width of tunnels formed by the colony (Pandya et al. 2022 ). On the other hand, feeding activity was evaluated from the quantity of food consumed and observations of trophallaxis among food-supplying ants. Foraging behaviour included the search, collection, and transport of food resources to the nest, while searching behaviour referred to ants moving within the setup, with or without antennal contact on the foraging ground. Immobility was noted in individuals that remained completely still, either isolated or aggregated away from nestmates. Communication was characterized through antennal contacts, body tapping, and trophallaxis between castes, whereas defensive behaviour was identified by immobile postures with raised antennae in response to perceived threats. Morphometric analysis of tunnel The tunneling pattern was assessed based on the number of burrows formed and the surface openings observed in the soil. Observations were recorded on days 7th ,14th, 21st, 28th, and 35th. Tunnel measurements were taken following the methodology outlined by Sinha ( 2014 ), wherein the number of openings, tunnel length, total depth, diameter, number of branches, and total burrow area were determined using the following formula: Area = π x a x b / 4 Where, a= length of burrow opening, b= width of burrow opening Estimation of neurotransmitters After a period of 7, 14, 21, 28 and 35 days, C. compressus individuals (n = 5) of media and minor each were sacrificed, brain was dissected and the levels of dopamine and serotonin were assessed according to Schlumfjf et al. ( 1974 ). Statistical analysis The experiments were performed in triplicate. One way analysis of variance (ANOVA) was used to compute the neurotransmitter levels in experimental worker minor and media, and their interaction, in comparison to controls. Post hoc comparisons were performed using Dunnett’s multiple comparison tests to determine the statistical significance ( p ≤ 0.05 ) and the analyses were done using the GraphPad Prism version 10. Results A notable abundance of C. compressus genus ant colonies in its locality of the site, the favourable environmental conditions, coupled with the visibly flourishing and active nature of these colonies, suggested that the ants were in a healthy state and thus ideal for the objectives of the study. Handpicking method offered valuable insights into the foraging behaviour and escape responses exhibited by ants across all caste types. Soil Analysis The collected soil was identified as sandy loam, which offered favourable conditions for ant nesting due to its low compaction. It showed a water holding capacity of 38.88%, and a soil porosity of 55.2% which is attributed to high drainage and the presence of large number of voids between the soil particles respectively. The soil had neutral pH of 7, which supports good nutrient availability. Behavioural studies During the acclimatization phase prior to queen removal, ants showed high responsiveness to food, characterized by rapid foraging, active communication between minor and media workers, minimal immobility, and continuous tunnel expansion, indicating favorable laboratory conditions. The removal of the queen had a significant effect on the behaviour of both minor and media worker ants of the experimental setup. Post the removal of the queen, these workers displayed noticeable reductions in communication, foraging, searching, feeding, and tunneling activities, alongside increase in immobility and defensive behaviours. These effects were most evident after 14 days (7 days post-removal). Over time, however, most behaviours gradually recovered, with many returning to near-normal levels by 28 days (Fig. 2). Searching behaviour was particularly affected, showing a marked decline in both workers, with minor as well as media experiencing a significant ( p < 0.05 ) reduction (Table 1 ). Additionally, an increase in size in one of the worker media ants was observed in the experimental colony after about 28 days of queen less environment which suggests the development of a plastic queen. Table 1 Chi square interactions between Minor and Media of C. compressus in laboratory Interactions X 2 p Communication Minor Media 7.63 2.77 0.106 0.597 Defense Minor Media 3.31 1.23 0.507 0.873 Feeding Minor Media 14.62 11.56 0.0056 0.021 Foraging Minor Media 3.18 2.67 0.527 0.61 Immobility Minor Media 11.45 6.37 0.02 0.17 Searching Minor Media 26.96 11.23 0.0001 0.02 Morphometric analysis of tunnel The removal of queen had a significant disturbance in the tunneling behaviour of the worker ants. A gradual and constant increase in the area and complexity of the tunnel was observed for the first 7 days in both the setups. After the removal of the queen from the experimental setup after 7 days, a breakdown of the tunnel structure was observed in causing the reduction in the area and complexity of the same. Gradually the tunneling behaviour started progressing and increasing gradually by 24th day (Table 2 ). The control group showed a constant increase of the tunneling pattern. Table 2 Tunneling pattern of C. compressus experimental setup Days Length (cm) Diameter (cm) Area (cm 2 ) Number of branches Type of tunneling 0–3 9.23 0.5 3.6 0 Simple 4–7 12.63 1.1 10.9 4 Complex 8–11 14.63 1.4 16.07 6 Complex 12–15 8.45 1.1 7.29 3 Complex 16–19 8.42 1.2 7.7 3 Complex 20–23 8.92 1.53 10.71 4 Complex 24–27 9.42 1.92 14.19 4 Complex 28–31 12.76 2.2 22.03 6 Complex 32–35 13.64 2.5 26.76 7 Complex Estimation of neurotransmitters A significant ( p < 0.05 ) decrease in the Dopamine and Serotonin levels in the brain tissues of worker media and minor was observed after 21 and 28 days post queen elimination in experimental setup in comparison to the constant high levels in the control group (Supplementary Table 1&2). The neurotransmitter levels of the workers of experimental setup gradually recover and become similar to that of the control group by 35 days (Fig. 3). Discussion Ants demonstrate impressive behavioural flexibility that goes well beyond simple stimulus-response patterns. This research emphasizes the importance of neuroplasticity in the brains of ants, particularly in relation to behavioural shifts as they move from performing tasks within the nest to foraging outside (Rössler 2019 ). The study centres on Camponotus compressus ants, which serve as a model species for investigating behavioural plasticity. Minor workers of C. compressus presented a significant variation in behaviour within 35 days of our study. The chosen site, having well balanced ecosystem indicated by the presence of thriving colonies functioning at their full potential, made it an ideal location for studying the natural behaviour of the ants and their environmental interactions, portraying an impact on the soil (pedological influence) occurring through the building of nests, tunnels, soil coverings, and mounds. These activities lead to changes in the soil's physical structure, as ants select, move, and reorganize soil particles (Bruyn and Conacher 1990 ; Cammeraat and Risch 2008 ). In our study, we found that, the highly porous nature and elevated water-holding capacity of sandy loam soil, coupled with a neutral pH created an ideal conditions for the nesting activity of C. compressus ants. As ground-dwelling insects that maintain direct contact with the substratum, these ants are particularly sensitive to soil characteristics. Previous studies suggests that sandy loam exhibits minimal compaction, which contributes to its high porosity and facilitates the construction of subterranean nest structures by the ants (Alaoui et al. 2017 ). Moreover, the neutral pH of 7 has been observed to support a diverse range of ant colonies, as higher soil pH levels are generally associated with increased species richness and thriving populations (Staab et al. 2014 ). Thus, our results are in accordance to the earlier reported studies. In our study, worker ant behaviours including foraging, feeding communication, searching as well as nesting were found to enhance tremendously during the first seven days in the presence of queen ant indicating a well-coordinated behavioural pattern within the colony. However, on the removal of the queen from the experimental colony led to significant behavioural changes among the worker ants. Overall, the findings suggest that queen presence plays a critical role in regulating worker activity, colony coordination, and behavioural stability, with partial behavioural compensation occurring over time in queenless colonies. Along with the modification in behaviour of worker ants, a significant change in the brain neurotransmitter levels for 35 days suggests correlation between neurotransmitter levels and behavioural activities. Results established that the absence of the queen disrupted the colony’s equilibrium, noticeably affecting key behavioural activities. It also led to measurable alterations in the levels of neurotransmitters, particularly dopamine and serotonin (Falibene et al. 2011 ; Friedman et al. 2018 ), in the worker ants. A relatively smaller decrease in the minor workers compared to the media workers attributes to the distinct roles each caste performs within the colony. Minor workers are primarily responsible for foraging and food collection and do not engage in direct communication with the queen. In contrast, media workers serve as intermediaries, directly interacting with the queen and relaying information throughout the colony (Jackson and Ratnieks 2006 ). As a result, the removal of the queen had a more profound impact on the media workers, leading to a greater decline in both neurotransmitter levels—such as dopamine and serotonin—and behavioural activities. The hierarchical nature of communication within the colony likely buffers the minor workers from the immediate effects of the queen's absence, as they primarily interact through the media caste. Thus, our findings suggest that the presence of the queen plays a crucial role in maintaining homeostasis within the colony, both socially and physiologically. Furthermore, by helping to regulate neurotransmitter levels, the queen ensures the proper functioning of worker ants, thereby supporting the overall well-being, coordination, and expansion of the colony and our work is in alignment to the previous studies (Falibene et al. 2011 ; Friedman et al. 2018 ; Aldana et al. 2024 ). Our study also reports the presence of an enlarged media worker in the experimental colony which indicated the emergence of a plastic queen, a worker ant that underwent physiological changes to assume the role of the queen in her absence. This phenomenon likely resulted from prolonged queenless environment, which triggered significant metabolic and hormonal shifts in certain workers. These changes can initiate the development of reproductive organs and lead to an increase in body size in a specific worker, eventually enabling her to take over the reproductive and regulatory functions of the colony. Once established, this plastic queen performs essential tasks such as mating, laying fertile eggs, and maintaining social order within the colony. Her presence helps restore stability, leading to normalized behavioural patterns and balanced neurotransmitter levels among the workers, allowing the colony to function efficiently once again (Friedman et al. 2018 ; Aldana et al. 2024 ). Declarations Acknowledgment The authors are truly grateful to the Department of Zoology, Faculty of Science, of The Maharaja Sayajirao University of Baroda for providing laboratory assistance. Authors are also thankful to the Department of Biochemistry for allowing the access of common instrumentation facility. Authors would also like to appreciate the assistance provided by summer interns-Yashvi, Rohit and Naksh in this research work. Competing Interest: The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Consent for Publication: Not applicable Funding: This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors. Data availability Statement: All data and materials used in the study are provided within this article or available from the corresponding author by request. References Adamo SA, Linn CE and Hoy RR (1995) The role of neurohormonal octopamine during ‘fight or flight’ behavior in the field cricket Gryllus bimaculatus . J. Exp. Biol. 198, 1691-1700. 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Current Opinion in Behavioral Sciences 39:98-105 . DOI: 10.1016/j.cobeha.2021.03.005 Sánchez-Tójar A, Moiron M, Niemela PT (2022) Terminology use in animal personality research: a self-report questionnaire and a systematic review. Proc. Roy. Soc. London. https://doi.org/10.1098/rspb.2021.2259 Saxena MM. 2001. Handbook of water and soil analysis. NPH books 169 pp. Scheiner R, Baumann A and Blenau W (2006) Aminergic control and modulation of honeybee behaviour. Curr. Neuropharmacol. 4 (4):259–276. http://dx.doi.org/10.2174/157015906778520791 Scheiner R, Entler BV, Barron AB, Scholl C and Thamm M (2017) The effects of fat body tyramine level on gustatory responsiveness of honeybees ( Apis mellifera ) differ between behavioral castes. 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Naturwissenschaften 92(4):198-201. https://doi.org/10.1007/s00114-005-0610-8 Sheikh AH,Iqbal J, Azad Z (2017) Study of Ant (Formicidae: Formicinae: Camponotus) fauna of Medical Hills Jabalpur, Madhya Pradesh. J. Adv. Res 3 (3), 44-46, 2017 Shimoji H, Aonuma H, Miura T, Tsuji K, Sasaki K and Okada Y (2017) Queen contact and among-worker interactions dually suppress worker brain dopamine as a potential regulator of reproduction in an ant. Behav Ecol Sociobiol. https://doi.org/10.1007/s00265-016-2263-3 Sih A, Bell AM and Johnson JC (2004a) Behavioral syndromes: an ecological and evolutionary overview. Trends Ecol Evol. . https://doi.org/10.1016/j.tree.2004.04.009 Sih A, Bell AM, Johnson JC and Ziemba RE (2004b) Behavioral syndromes: an integrative overview. Q. Rev. Biol . https://doi.org/10.1086/422893 Simpson SJ and Raubenheimer D (2012) The Nature of Nutrition: A Unifying Framework from Animal Adaptation to Human Obesity. Princeton University Press (book). ISBN: 9780691145658 Sinha S (2014) Morphological characteristics and distribution of the burrows of freshwater crab Barytelphusa cunicularis (Westwood, 1836). Indian J. Sci. Res 4: 158-164. Smith, Kamhi JF, Traniello JF (2013) Biogenic amines and collective organization in a superorganism: neuromodulation of social behavior in ants. Brain Behav. Evol. 82(4):220-36. https://doi.org/10.1159/000356091 Soares PAO, Delabie JHC, Zanuncio JCand Serrão JE (2008) Neural Plasticity in The Brain of Workers of the Carpenter Ant Camponotus rufipes (Hymenoptera: Formicidae). Sociobiology 51 (3), 705, 2008 Staab M, Schuldt A, Assmann T and Klein AM (2014) Tree diversity promotes predator but not omnivore ants in a subtropical Chinese forest. Ecol. Entomol. 39 :637–647. http://dx.doi.org/10.1111/een.12143 Steck K, Hansson BS and Knaden M (2009) Smells like home: Desert ants, Cataglyphis fortis , use olfactory landmarks to pinpoint the nest. Front. Zool. doi:10.1186/1742-9994-6-5 Stevenson PA, Dyakonova V, Rillich J and Schildberger K (2005) Octopamine and experience-dependent modulation of aggression in crickets. J. Neurosci. 25, 1431-1441 Stone AM, Terblanche JS, Pottier J, Kellermann V, and Chown SL (2022) Meta-analysis reveals weak but pervasive plasticity in insect thermal limits. Nat. Commun. 13 , 5292. http://dx.doi.org/10.1038/s41467-022-32953-2 Szabó N, Nagy J and Tartally A (2025) Comparison of the simplest diets to find the most effective one in developing ant colonies of Lasius niger . Sci. Rep. 15 :20540. http://dx.doi.org/10.1038/s41598-025-06406-x Szczuka A, Korczyńska J, Wnuk A, Symonowicz B, Gonzalez-Szwacka A, Mazurkiewicz P, Kostowski W and Godzińska EJ (2013) The effects of serotonin, dopamine, octopamine and tyramine on behavior of workers of the ant Formica polyctena during dyadic aggression tests. Acta Neurobiol. Exp. (Warsaw). https://doi.org/10.55782/ane-2013-1955 Toth AL and Dolezal AG (2017) Molecular evolution of insect sociality: an eco-evo-devo perspective. Annu. Rev. Entomol. 62 :419–442. http://dx.doi.org/10.1146/annurev-ento-031616-035601 Trible W and Kronauer DJC (2017) A caste differentiation mutant elucidates the evolution of ant caste development. J Exp Biol. https://doi.org/10.1242/jeb.145292 Tschinkel WR (2004) The nest architecture of the Florida harvester ant, Pogonomyrmex badius. J. Insect Sci. Volume 4, Issue 1. https://doi.org/10.1093/jis/4.1.21 Vander Meer RK, Morel L (2008) Queen regulates biogenic amine level and nestmate recognition in workers of the fire ant Solenopsis invicta . Naturwissenschaften. https://doi.org/10.1007/s00114-008-0432-6 Witte V, Attygalle AB and Meinwald J (2007) Complex chemical communication in the crazy ant Paratrechina longicornis Latreille (Hymenoptera: Formicidae). Chemoecology. https://doi.org/10.1007/s00049-006-0364-6 Wystrach A and Graham P (2012) What can we learn from studies of insect navigation? Anim. Behav. Vol 84, No. 1. https://doi.org/10.1016/j.anbehav.2012.04.017 Supplementary Files Supplementaryfile.docx Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-8751427","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":586541298,"identity":"51498f76-c1c5-4774-88af-79cd6db661ac","order_by":0,"name":"Swasti Saxena","email":"","orcid":"","institution":"The Maharaja Sayajirao University of Baroda","correspondingAuthor":false,"prefix":"","firstName":"Swasti","middleName":"","lastName":"Saxena","suffix":""},{"id":586541299,"identity":"62001164-b496-48e1-af56-42b85f1b5039","order_by":1,"name":"Ankita Salunke","email":"","orcid":"","institution":"The Maharaja Sayajirao University of Baroda","correspondingAuthor":false,"prefix":"","firstName":"Ankita","middleName":"","lastName":"Salunke","suffix":""},{"id":586541300,"identity":"4b48a9aa-cee9-4650-ba36-e4cb546bb04b","order_by":2,"name":"Nishi Pandya","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA90lEQVRIiWNgGAWjYDACZgY2hgQQ4wAD+4cPQJqNnQQtbIwzQFqYCdvDBqGAWph5IIbgB+bs7M8ePNxhl8d3I/nZY5tf2+T5mBkYP3zMwa3FspnH3CDxTHKx5I00c+PcvtuGbcwMzJIzt+HWYnCYh00isY05ccONBAPp3J7bjEAtbMy8eLWwPwNqqQdqSf8gbdlz254ILQxmQC2HgVpyzKQZftxOJEILD1DLmeOJM8+8KTbsbbid3MbM2IzfL+ePP5P8uaM6se94+sYHP/7ctp3f3nzww0c8WsCAsQFICCQAGW1wLjFa+A8AiT+EFY+CUTAKRsHIAwBujFWbKn8QCwAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0009-0004-0552-2286","institution":"The Maharaja Sayajirao University of Baroda","correspondingAuthor":true,"prefix":"","firstName":"Nishi","middleName":"","lastName":"Pandya","suffix":""}],"badges":[],"createdAt":"2026-01-31 16:51:35","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8751427/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8751427/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":102345988,"identity":"4929af81-911d-4e19-8059-8cc4794ea9be","added_by":"auto","created_at":"2026-02-10 17:30:47","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":39028,"visible":true,"origin":"","legend":"\u003cp\u003eLegend not included with this version\u003c/p\u003e","description":"","filename":"Fig1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8751427/v1/7d2c2c6a9619ec3c75f4a009.jpg"},{"id":102345987,"identity":"0300bf72-a7f4-4baf-b2a1-1318fb85c8b5","added_by":"auto","created_at":"2026-02-10 17:30:47","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":233074,"visible":true,"origin":"","legend":"\u003cp\u003eLegend not included with this version\u003c/p\u003e","description":"","filename":"Fig2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8751427/v1/cd06b3945c4e061f563faf23.jpg"},{"id":102345990,"identity":"71dd9603-27f2-4ca5-b9a1-0349643ff4ec","added_by":"auto","created_at":"2026-02-10 17:30:47","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":111517,"visible":true,"origin":"","legend":"\u003cp\u003eLegend not included with this version\u003c/p\u003e","description":"","filename":"Fig3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8751427/v1/295b105a7fd62915e41abe85.jpg"},{"id":106727005,"identity":"801c25ea-1536-432d-9222-43765213d4bb","added_by":"auto","created_at":"2026-04-12 18:37:58","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1634321,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8751427/v1/76dfd73e-f435-4f4d-acff-2e93eb34c095.pdf"},{"id":102345989,"identity":"81f6da13-3449-490f-b316-4ce1d7abe65f","added_by":"auto","created_at":"2026-02-10 17:30:47","extension":"docx","order_by":8,"title":"","display":"","copyAsset":false,"role":"supplement","size":15268,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementaryfile.docx","url":"https://assets-eu.researchsquare.com/files/rs-8751427/v1/92ee18a04e03ca20bf1c138d.docx"}],"financialInterests":"","formattedTitle":"Elucidating the role of neurotransmitters in the behavioural plasticity of Camponotus compressus (Hymenoptera: Formicidae)","fulltext":[{"header":"Introduction","content":"\u003cp\u003eAnimal personality and behavioural syndromes, conflated in earlier literature (MacKay and Haskell \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; S\u0026aacute;nchez-T\u0026oacute;jar et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), represents distinct conceptual frameworks: animal personality describes individual consistency in specific traits across contexts including aggression stability across ecological fluctuations (Bell and Sih \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), while behavioural syndromes like boldness-aggressiveness and associations address traits correlations within individuals (Sih et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2004a\u003c/span\u003e, \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003eb\u003c/span\u003e). Some studies have demonstrated the absence of significant correlations between behavioural traits (Carvalho et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Roth et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Insects, frequently regarded as less adaptable than vertebrates (Chittka and Niven \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Wystrach and Graham \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), typically exhibit compensatory abilities for impairments as immediate responses, attributed to the inherent natural redundancy and robustness of their systems, rather than as outcomes of lifelong neural plasticity (Wystrach and Graham \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Stone et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Insects like ants, cockroaches, stick insects and other insects, for instance, show a robust and spontaneous neural response governing leg coordination (Steck et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Schwarz et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eInsect colonies are formed of a number of individuals each representing a distinct labour such as reproductive division of tasks between queen, king (in termites), largely sterile workers as well as communication between and within the caste for the efficient functioning of a colony, essential for ecological success (Hӧlldobler and Wilson 1990). Workers respond to the changing signals in a way that are beneficial for the entire colony (Hӧlldobler and Wilson 2009).\u003c/p\u003e \u003cp\u003eAnts, including largely of castes\u0026mdash;queen, drone (male), and worker ants (Clark and Holbrook \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) exhibit a distinctive nesting pattern characterised by social interaction in nest construction. Communication within the colony adheres to a hierarchical framework, wherein minor ants predominantly engage with medium ants, who subsequently convey information to the queen (Wollman et al. 2011). Consequently, they construct a nest featuring intricate, network-like structures composed of chambers interconnected by tunnels that serve as nodes and connecting links, respectively (Tschinkel \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Buhl et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Improved communication among chambers can accelerate resource transfer inside the nest (Cook et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eMembers of a colony are divided into castes with distinct characteristics, such as behaviour, metabolism, lifespan, and reproduction. Only a single queen is accountable for reproduction, possessing ovarioles with oocytes at various stages of differentiation (Khila and Abouheif \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Lee et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Penick et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Conversely, workers are non-reproductive, possessing undeveloped ovaries which constitute that exhibit distinct characteristics from the queen, engaging in incessant movement, brood care, colony sanitation, and food foraging, which are acquired during the embryonic and larval developmental stages (Khila and Abouheif \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2010\u003c/span\u003e: Schwander et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Trible and Kronauer 2021). Queen signalling is crucial for ant colonies as it signals the queen's presence and reproductive status, prompting workers to forgo their own reproductive efforts and focus on assisting with brood care (Keller and Nonacs \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Leonhardt et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Moreover, worker ants signal and encourage others to follow them to the food source through tandem running (Witte et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Holldobler and Wilson 2009; Simpson and Raubenheimer \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Csata and Dussutour \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFemale workers typically do not reproduce; instead, they are responsible for colony maintenance and brood care. However, despite this clear division of roles, workers can become reproductively active under certain conditions. The presence of a queen strongly suppresses this reproductive flexibility. In queenless species, colony reproduction is managed through a well-organized system where reproductive and non-reproductive tasks are distinctly divided among workers. The reproductive ability in a queenless colony lies to some individuals of the colony, for example, in \u003cem\u003eNeoponera apicalis\u003c/em\u003e, the workers establish a hierarchy based on the behaviours such as antennal dueling and biting. One of the dominant higher worker which exhibits a higher frequency of this behaviour, lays most of the eggs which result into male progenies. Also, in \u003cem\u003eHarpegnathos saltator\u003c/em\u003e, the colony is controlled by one queen or a bunch of gamergates which is a mated worker, with a sexually reproducing spermatheca (H\u0026ouml;lldobler and Wilson \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Peeters and H\u0026ouml;lldobler \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Monnin and Peeters \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Aldana et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eInterpreting the concept of regulation in social insects poses a distinct challenge, especially in comprehending the neurological mechanisms that link behavioural conditioning in the brain to alterations in reproductive physiology. Biogenic amines act as neurotransmitters, neuromodulators, and neurohormones in the brain, playing a vital role in the regulation of behaviour (Roeder \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Scheiner et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Pandya et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In insects, the competition for reproductive hierarchy occurs among the colony\u0026rsquo;s females with behavioural consequences frequently regulated by neurochemical pathways. Moreover, the concentration of biogenic amines in brain tissue can profoundly influence individual worker behaviour by modulating their responsiveness to environmental inputs, including foraging (Friedman et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Serotonin and dopamine, as biogenic amines, are recognized for their role in regulating dominance-related behaviours, particularly aggression among colony caste members (Miczek et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Kravitz and Huber \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Nelson \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Octopamine and serotonin are associated with aggression in crickets (Adamo et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Murakami and Itoh \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Dyakonova et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Barbero et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2023\u003c/span\u003e); dopamine has been linked to social dominance in both vertebrates and solitary insects (Baier et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Miczek et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Stevenson et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Biogenic amines regulate key aspects of ant behaviour, with serotonin and dopamine modulating aggression, defense, and arousal, while octopamine and tyramine influence social interactions and aggression control. Additionally, shifts in amino acid ratios with age and task specialization underscore their role in coordinating both individual and collective colony functions (Seid and Traniello \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Smith et al. 2013; Szczuka 2013; Koyama et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Aonuma \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBiogenic amines modulates aggression in \u003cem\u003eFormica polyctena\u003c/em\u003e ants, wherein exogenous administration of serotonin, dopamine, octopamine, and tyramine was assessed during dual aggression tests involving nest mates, alien conspecifics, allospecific ants (\u003cem\u003eFormica fusca\u003c/em\u003e), and prey (house cricket nymphs, \u003cem\u003eAcheta domesticus\u003c/em\u003e) (Vander Meer et al. 2008; Ohkawara and Aonuma \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Shimoji et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Serotonin gradually enhances physical aggression, encompassing biting and the secretion of formic acid whereas dopamine increases the mandible threats and biting toward both \u003cem\u003eF. fusca\u003c/em\u003e and cricket nymphs (Szczuka et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). On the other hand, octopamine exhibited no substantial impact on aggressive behaviours. Such variations in reproductive status and behaviour among nestmates in ant and bee species are further associated with tissue-specific physiological and transcriptomic differences (Jandt and Gordon \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Jeanne \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Toth and Dolezal \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Chandra et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In particular, biogenic amine and neurohormonal signaling in the brains of workers are crucial for regulating foraging behaviour in both social and solitary insects. Alterations in brain biogenic amine concentrations can influence individual workers\u0026rsquo; responses to specific stimuli, including foraging cues (Muscedere et al. 2012; Kamhi and Traniello \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Bubak et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Scheiner et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Natural differences in nestmates' sensitivity to stimuli can be beneficial for colony function, such as enabling flexible task allocation.\u003c/p\u003e \u003cp\u003eAnts are considered a highly suitable model organism for exploring the neurobiological studies of foraging behaviour of group animals to account the highly organized structures built due to the collective efforts of the colony (Li et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Studies suggest that elimination of queen may cause imbalance in the neurotransmitter levels and behavioural activities which might result into neural plasticity. However, there is a lacuna in the studies conducted on behavioural plasticity of commonly found ants like the \u003cem\u003eC. compressus\u003c/em\u003e. Therefore, the present study was aimed to elucidate the role of neurotransmitters in plasticity of \u003cem\u003eC. compressus\u003c/em\u003e ants. We assessed the behavioural plasticity in worker ants by evaluating the nesting behaviour, levels of neurotransmitters and whether a worker ant achieves the plasticity or not.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eCollection, rearing and acclimatisation\u003c/h2\u003e \u003cp\u003e \u003cem\u003eCamponotus compressus\u003c/em\u003e were collected from the grounds of the Pavilion of Maharaja Sayajirao University of Baroda (22.320153\u003csup\u003eo\u003c/sup\u003eN, 73.185734\u003csup\u003eo\u003c/sup\u003eE and 22.31992\u003csup\u003eo\u003c/sup\u003eN, 73.185851\u003csup\u003eo\u003c/sup\u003eE) of Vadodara city, India. Collection of \u003cem\u003eC. compressus\u003c/em\u003e was carried out during the noon in the months of December 2024 to May 2025, using handpicking method from the ground and bark of the tree. The collected ant colonies were brought to laboratory for identification and rearing. Morphological identification was done up to the species level with the help of standard taxonomic keys and by comparing the specimens in department repository (Sheikh et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The ants\u0026minus;30 minor, 19 media, and 1 queen were reared in nesting and foraging boxes (3 sets, sized-15.0 \u0026sdot; 8.5 \u0026sdot; 20 cm) under laboratory conditions (26\u003csup\u003e\u0026deg;\u003c/sup\u003eC temperature, 75% humidity and 12h/12h night and day cycles), using the method of Soares et al. (\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) and Heinze \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2020\u003c/span\u003e (Fig.\u0026nbsp;1). They were fed with supplements like sugar solution (4mL of 40% solution) for the carbohydrate content while freshly dead or alive but crushed insects for the protein requirement of the ants (Szab\u0026oacute; et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The dead ants were removed from the setup to eliminate any negative effects on the rearing and maintenance.\u003c/p\u003e \u003cp\u003eThe soil required for the rearing was collected from the same sampling sites at the time of ant collection in order to provide a suitable environment (Czechowski and Pisarski \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1992\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eSoil Analysis\u003c/h3\u003e\n\u003cp\u003eThe physical and chemical parameters of the collected soil were checked to analyse the type of soil and its quality in relation to \u003cem\u003eC. compressus.\u003c/em\u003e Soil texture identification was done by texture analysis and further, the soil porosity, water holding capacity and pH were measured using the method given by Saxena (\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2001\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eBehavioural studies\u003c/h3\u003e\n\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eExperimental setup\u003c/h2\u003e \u003cp\u003eAfter acclimatisation period of the ants in the control and experimental setup (30 minor, 19 media and 1 queen for each group), the behavioural activities of \u003cem\u003eC. compressus\u003c/em\u003e were observed and recorded on 7th ,14th, 21st, 28th, and 35th day. In the experimental setup, the queen ant was removed on the 7th day and the worker ants were observed for their behaviour and neural regulation in absence of queen. Food was provided at regular intervals to ensure the normal functioning and behaviour of \u003cem\u003eC. compressus\u003c/em\u003e.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eBehaviour\u003c/h3\u003e\n\u003cp\u003eSeveral behavioural patterns such as tunneling, feeding, foraging, searching, immobility, communication, and defence were examined across the experimental and control group. For each behaviour, readings were taken after every four days, with each session lasting five hours comprising of one-hour intervals between every observation. The ant colonies were observed regularly to analyze acclimatisation to the laboratory conditions.\u003c/p\u003e \u003cp\u003eThe behavioural reservoire of \u003cem\u003eC. compressus\u003c/em\u003e under laboratory conditions was assessed through multiple parameters. Tunneling behaviour was recorded by measuring the number, length, and width of tunnels formed by the colony (Pandya et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). On the other hand, feeding activity was evaluated from the quantity of food consumed and observations of trophallaxis among food-supplying ants. Foraging behaviour included the search, collection, and transport of food resources to the nest, while searching behaviour referred to ants moving within the setup, with or without antennal contact on the foraging ground. Immobility was noted in individuals that remained completely still, either isolated or aggregated away from nestmates. Communication was characterized through antennal contacts, body tapping, and trophallaxis between castes, whereas defensive behaviour was identified by immobile postures with raised antennae in response to perceived threats.\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eMorphometric analysis of tunnel\u003c/h2\u003e \u003cp\u003eThe tunneling pattern was assessed based on the number of burrows formed and the surface openings observed in the soil. Observations were recorded on days 7th ,14th, 21st, 28th, and 35th. Tunnel measurements were taken following the methodology outlined by Sinha (\u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), wherein the number of openings, tunnel length, total depth, diameter, number of branches, and total burrow area were determined using the following formula:\u003c/p\u003e \u003cp\u003eArea\u0026thinsp;=\u0026thinsp;π x a x b / 4\u003c/p\u003e \u003cp\u003eWhere, a= length of burrow opening, b= width of burrow opening\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eEstimation of neurotransmitters\u003c/h3\u003e\n\u003cp\u003eAfter a period of 7, 14, 21, 28 and 35 days, \u003cem\u003eC. compressus\u003c/em\u003e individuals (n\u0026thinsp;=\u0026thinsp;5) of media and minor each were sacrificed, brain was dissected and the levels of dopamine and serotonin were assessed according to Schlumfjf et al. (\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e1974\u003c/span\u003e).\u003c/p\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eThe experiments were performed in triplicate. One way analysis of variance (ANOVA) was used to compute the neurotransmitter levels in experimental worker minor and media, and their interaction, in comparison to controls. Post hoc comparisons were performed using Dunnett\u0026rsquo;s multiple comparison tests to determine the statistical significance (\u003cem\u003ep \u0026le; 0.05\u003c/em\u003e) and the analyses were done using the GraphPad Prism version 10.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eA notable abundance of \u003cem\u003eC. compressus\u003c/em\u003e genus ant colonies in its locality of the site, the favourable environmental conditions, coupled with the visibly flourishing and active nature of these colonies, suggested that the ants were in a healthy state and thus ideal for the objectives of the study. Handpicking method offered valuable insights into the foraging behaviour and escape responses exhibited by ants across all caste types.\u003c/p\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eSoil Analysis\u003c/h2\u003e \u003cp\u003eThe collected soil was identified as sandy loam, which offered favourable conditions for ant nesting due to its low compaction. It showed a water holding capacity of 38.88%, and a soil porosity of 55.2% which is attributed to high drainage and the presence of large number of voids between the soil particles respectively. The soil had neutral pH of 7, which supports good nutrient availability.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eBehavioural studies\u003c/h2\u003e \u003cp\u003eDuring the acclimatization phase prior to queen removal, ants showed high responsiveness to food, characterized by rapid foraging, active communication between minor and media workers, minimal immobility, and continuous tunnel expansion, indicating favorable laboratory conditions.\u003c/p\u003e \u003cp\u003eThe removal of the queen had a significant effect on the behaviour of both minor and media worker ants of the experimental setup. Post the removal of the queen, these workers displayed noticeable reductions in communication, foraging, searching, feeding, and tunneling activities, alongside increase in immobility and defensive behaviours. These effects were most evident after 14 days (7 days post-removal). Over time, however, most behaviours gradually recovered, with many returning to near-normal levels by 28 days (Fig.\u0026nbsp;2). Searching behaviour was particularly affected, showing a marked decline in both workers, with minor as well as media experiencing a significant (\u003cem\u003ep\u0026thinsp;\u0026lt;\u0026thinsp;0.05\u003c/em\u003e) reduction (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Additionally, an increase in size in one of the worker media ants was observed in the experimental colony after about 28 days of queen less environment which suggests the development of a plastic queen.\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\u003eChi square interactions between Minor and Media of \u003cem\u003eC. compressus\u003c/em\u003e in laboratory\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"3\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInteractions\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eX\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003ep\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCommunication\u003c/p\u003e \u003cp\u003eMinor\u003c/p\u003e \u003cp\u003eMedia\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7.63\u003c/p\u003e \u003cp\u003e2.77\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.106\u003c/p\u003e \u003cp\u003e0.597\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDefense\u003c/p\u003e \u003cp\u003eMinor\u003c/p\u003e \u003cp\u003eMedia\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3.31\u003c/p\u003e \u003cp\u003e1.23\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.507\u003c/p\u003e \u003cp\u003e0.873\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eFeeding\u003c/b\u003e\u003c/p\u003e \u003cp\u003eMinor\u003c/p\u003e \u003cp\u003eMedia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14.62\u003c/p\u003e \u003cp\u003e11.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.0056\u003c/p\u003e \u003cp\u003e0.021\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eForaging\u003c/b\u003e\u003c/p\u003e \u003cp\u003eMinor\u003c/p\u003e \u003cp\u003eMedia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3.18\u003c/p\u003e \u003cp\u003e2.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.527\u003c/p\u003e \u003cp\u003e0.61\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eImmobility\u003c/b\u003e\u003c/p\u003e \u003cp\u003eMinor\u003c/p\u003e \u003cp\u003eMedia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e11.45\u003c/p\u003e \u003cp\u003e6.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.02\u003c/p\u003e \u003cp\u003e0.17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSearching\u003c/b\u003e\u003c/p\u003e \u003cp\u003eMinor\u003c/p\u003e \u003cp\u003eMedia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e26.96\u003c/p\u003e \u003cp\u003e11.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.0001\u003c/p\u003e \u003cp\u003e0.02\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eMorphometric analysis of tunnel\u003c/h2\u003e \u003cp\u003eThe removal of queen had a significant disturbance in the tunneling behaviour of the worker ants. A gradual and constant increase in the area and complexity of the tunnel was observed for the first 7 days in both the setups. After the removal of the queen from the experimental setup after 7 days, a breakdown of the tunnel structure was observed in causing the reduction in the area and complexity of the same. Gradually the tunneling behaviour started progressing and increasing gradually by 24th day (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The control group showed a constant increase of the tunneling pattern.\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\u003eTunneling pattern of \u003cem\u003eC. compressus\u003c/em\u003e experimental setup\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDays\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLength (cm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDiameter (cm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eArea (cm\u003csup\u003e2\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNumber of branches\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eType of tunneling\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e0\u0026ndash;3\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e9.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e3.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSimple\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e4\u0026ndash;7\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e12.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eComplex\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e8\u0026ndash;11\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e14.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e16.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eComplex\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e12\u0026ndash;15\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eComplex\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e16\u0026ndash;19\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eComplex\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e20\u0026ndash;23\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1.53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10.71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eComplex\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e24\u0026ndash;27\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e9.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e14.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eComplex\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e28\u0026ndash;31\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e12.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e22.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eComplex\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e32\u0026ndash;35\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e13.64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e26.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eComplex\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eEstimation of neurotransmitters\u003c/h2\u003e \u003cp\u003eA significant (\u003cem\u003ep\u0026thinsp;\u0026lt;\u0026thinsp;0.05\u003c/em\u003e) decrease in the Dopamine and Serotonin levels in the brain tissues of worker media and minor was observed after 21 and 28 days post queen elimination in experimental setup in comparison to the constant high levels in the control group (Supplementary Table\u0026nbsp;1\u0026amp;2). The neurotransmitter levels of the workers of experimental setup gradually recover and become similar to that of the control group by 35 days (Fig.\u0026nbsp;3).\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eAnts demonstrate impressive behavioural flexibility that goes well beyond simple stimulus-response patterns. This research emphasizes the importance of neuroplasticity in the brains of ants, particularly in relation to behavioural shifts as they move from performing tasks within the nest to foraging outside (R\u0026ouml;ssler \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The study centres on \u003cem\u003eCamponotus compressus\u003c/em\u003e ants, which serve as a model species for investigating behavioural plasticity. Minor workers of \u003cem\u003eC. compressus\u003c/em\u003e presented a significant variation in behaviour within 35 days of our study.\u003c/p\u003e \u003cp\u003eThe chosen site, having well balanced ecosystem indicated by the presence of thriving colonies functioning at their full potential, made it an ideal location for studying the natural behaviour of the ants and their environmental interactions, portraying an impact on the soil (pedological influence) occurring through the building of nests, tunnels, soil coverings, and mounds. These activities lead to changes in the soil's physical structure, as ants select, move, and reorganize soil particles (Bruyn and Conacher \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Cammeraat and Risch \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). In our study, we found that, the highly porous nature and elevated water-holding capacity of sandy loam soil, coupled with a neutral pH created an ideal conditions for the nesting activity of \u003cem\u003eC. compressus\u003c/em\u003e ants. As ground-dwelling insects that maintain direct contact with the substratum, these ants are particularly sensitive to soil characteristics. Previous studies suggests that sandy loam exhibits minimal compaction, which contributes to its high porosity and facilitates the construction of subterranean nest structures by the ants (Alaoui et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Moreover, the neutral pH of 7 has been observed to support a diverse range of ant colonies, as higher soil pH levels are generally associated with increased species richness and thriving populations (Staab et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Thus, our results are in accordance to the earlier reported studies.\u003c/p\u003e \u003cp\u003eIn our study, worker ant behaviours including foraging, feeding communication, searching as well as nesting were found to enhance tremendously during the first seven days in the presence of queen ant indicating a well-coordinated behavioural pattern within the colony. However, on the removal of the queen from the experimental colony led to significant behavioural changes among the worker ants. Overall, the findings suggest that queen presence plays a critical role in regulating worker activity, colony coordination, and behavioural stability, with partial behavioural compensation occurring over time in queenless colonies. Along with the modification in behaviour of worker ants, a significant change in the brain neurotransmitter levels for 35 days suggests correlation between neurotransmitter levels and behavioural activities. Results established that the absence of the queen disrupted the colony\u0026rsquo;s equilibrium, noticeably affecting key behavioural activities. It also led to measurable alterations in the levels of neurotransmitters, particularly dopamine and serotonin (Falibene et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Friedman et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), in the worker ants. A relatively smaller decrease in the minor workers compared to the media workers attributes to the distinct roles each caste performs within the colony. Minor workers are primarily responsible for foraging and food collection and do not engage in direct communication with the queen. In contrast, media workers serve as intermediaries, directly interacting with the queen and relaying information throughout the colony (Jackson and Ratnieks \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). As a result, the removal of the queen had a more profound impact on the media workers, leading to a greater decline in both neurotransmitter levels\u0026mdash;such as dopamine and serotonin\u0026mdash;and behavioural activities. The hierarchical nature of communication within the colony likely buffers the minor workers from the immediate effects of the queen's absence, as they primarily interact through the media caste. Thus, our findings suggest that the presence of the queen plays a crucial role in maintaining homeostasis within the colony, both socially and physiologically. Furthermore, by helping to regulate neurotransmitter levels, the queen ensures the proper functioning of worker ants, thereby supporting the overall well-being, coordination, and expansion of the colony and our work is in alignment to the previous studies (Falibene et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Friedman et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Aldana et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOur study also reports the presence of an enlarged media worker in the experimental colony which indicated the emergence of a plastic queen, a worker ant that underwent physiological changes to assume the role of the queen in her absence. This phenomenon likely resulted from prolonged queenless environment, which triggered significant metabolic and hormonal shifts in certain workers. These changes can initiate the development of reproductive organs and lead to an increase in body size in a specific worker, eventually enabling her to take over the reproductive and regulatory functions of the colony. Once established, this plastic queen performs essential tasks such as mating, laying fertile eggs, and maintaining social order within the colony. Her presence helps restore stability, leading to normalized behavioural patterns and balanced neurotransmitter levels among the workers, allowing the colony to function efficiently once again (Friedman et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Aldana et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgment\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors are truly grateful to the Department of Zoology, Faculty of Science, of The Maharaja Sayajirao University of Baroda for providing laboratory assistance. Authors are also thankful to the Department of Biochemistry for allowing the access of common instrumentation facility. Authors would also like to appreciate the assistance provided by summer interns-Yashvi, Rohit and Naksh in this research work.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting Interest:\u0026nbsp;\u003c/strong\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for Publication:\u003c/strong\u003e Not applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding:\u003c/strong\u003e This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability Statement:\u003c/strong\u003e All data and materials used in the study are provided within this article or available from the corresponding author by request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAdamo SA, Linn CE and Hoy RR (1995) The role of neurohormonal octopamine during \u0026lsquo;fight or flight\u0026rsquo; behavior in the field cricket \u003cem\u003eGryllus bimaculatus\u003c/em\u003e. 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Ecol. 24(5): 1081\u0026ndash;1091.https://doi.org/10.1093/beheco/art034\u003c/li\u003e\n\u003cli\u003e\u003cstrong\u003eChandra V, Fetter-Pruneda I, Oxley PR, Ritger AL, McKenzie SK, Libbrecht R and Kronauer DJC (2018)\u003c/strong\u003e Social regulation of insulin signaling and the evolution of eusociality in ants. \u003cem\u003eScience\u003c/em\u003e. https://doi.org/10.1126/science.aar5723\u003c/li\u003e\n\u003cli\u003eChittka L and Niven J (2009) Are Bigger Brains Better? Curr Biol Vol 19 No 21. DOI: 10.1016/j.cub.2009.08.023\u003c/li\u003e\n\u003cli\u003eClark R and Holbrook T (2009) Ant Farm- Ask a Biologist activity for classroom and home.\u003c/li\u003e\n\u003cli\u003eCook Z, Franks DW, Robinson EJH (2014) Efficiency and robustness of ant colony transportation networks. Behav. Ecol. 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Vol 84, No. 1. https://doi.org/10.1016/j.anbehav.2012.04.017\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Camponotus compressus, Nesting, Behaviour, Neuroplasticity, Dopamine, Serotonin","lastPublishedDoi":"10.21203/rs.3.rs-8751427/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8751427/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eAnts express a remarkable behavioural plasticity ranging far beyond rigid stimulus-response relationships. They are omnivorous and each caste exhibit a variety of behaviour including nesting (foraging, food storage, protection from predators, feeding, nest structure and brood care). The behaviour investment of workers is shaped by presence or absence of queen within the colony; which may affect the behavioural flexibility and developmental neuroplasticity in workers. However, there is a lacuna in the studies conducted in the nesting behaviour and the associated neurophysiology of such ants. So, the present study was aimed to determine the role of neurotransmitters (Dopamine and Serotonin) underlining neuroplasticity in the nesting behaviour\u0026minus;foraging, feeding, searching, immobility, communication, defence, and nest structure of \u003cem\u003eCamponotus compressus\u003c/em\u003e ant, under laboratory conditions. Our results demonstrated an increase in the respective behavioural activities, as well as in the length and depth of the nest, in the presence of the queen, with a subsequent decrease on the 14th day following her removal. Additionally, neurotransmitter levels (DA and 5-HT) were higher in workers (media and minor) ants on the 7th day, followed by a significant (\u003cem\u003ep\u0026thinsp;\u0026lt;\u0026thinsp;0.05\u003c/em\u003e) decline after 14th day suggesting a reduced nesting activity, which is further correlated with the decrease in nest morphometry, in comparison to control. The current study unveils the neurophysiological mechanism involved in achieving neuroplasticity in the \u003cem\u003eC. compressus\u003c/em\u003e. Further studies on specific gene expressions and histochemistry of brain will provide a broader view on the altered nesting behaviour in worker ants and the developed plastic queen.\u003c/p\u003e","manuscriptTitle":"Elucidating the role of neurotransmitters in the behavioural plasticity of Camponotus compressus (Hymenoptera: Formicidae)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-02-10 17:30:43","doi":"10.21203/rs.3.rs-8751427/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"c9df1859-2e46-47aa-b64f-6be2a20b6980","owner":[],"postedDate":"February 10th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-04-10T22:16:04+00:00","versionOfRecord":[],"versionCreatedAt":"2026-02-10 17:30:43","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8751427","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8751427","identity":"rs-8751427","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}