Core-periphery structure of a medicinal botanical system in Uruguay

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These experiences manifest into cultural traits, influencing human culture, particularly in healthcare and maintenance. Studies regarding medicinal plants and treatments are integral to the study of the medical botanical system. Pharmacopeias highlight the prevalence of specific species widely used, aligning with the “consensus within diversity theory” in evolutionary ethnobiology. Methods: This study applies network analysis on the botanical system of “Parque Regional Quebradas del Norte”, Rivera, Uruguay. Results: Study results demonstrate a core-periphery structure, with a tightly interconnected core resistant to fragmentation, ensuring structural stability. This study showed the existence of peripheral nodes in the entire system, improving resilience against disturbances and increasing population's health. Conclusion: The core species, renowned for their versatility and multiple medicinal uses, treating less severe ailments effectively. Additionally, core plants serve as prototypes for innovations. Their extinction poses a threat to the system's resilience. Conversely, peripheral plants, though vulnerable, offer possibilities for therapeutic innovations. In the face of environmental change, conservation efforts should prioritize species that are vulnerable to extinction, particularly within the core. Simultaneously, preserving knowledge associated with peripheral plants presents a bicultural conservation strategy, ensuring the botanical system's robustness amongst evolving ecological conditions. Adaptive memory Core-periphery structure Medical Botanical System Medicinal plants Network analysis Resilience Traditional knowledge Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Throughout the course of adaptation, humans have developed cognitive processes to confront the difficulties presented by nature. As a result, natural selection has played a crucial role in shaping individuals' psychological structure, enabling them to remember experiences and enhance their ability to survive. Sensations and perceptions are naturally interpreted to create a response to difficult and stimulating environments [ 1 – 4 ]. Nairne & Pandeirada [ 3 ] propose that experiences are generationally remembered, integrated into an adaptive memory system. It has been documented that cultural features respond to situations of risk, partner preference, illness prevention, healing processes and illness therapies [ 5 – 9 ]. These cultural traits are transmitted within a population through various pathways or mechanisms of knowledge diffusion: a) from parents to children (vertical); b) among peers of the same generation (horizontal); c) between generations, excluding the parenteral route (oblique); d) from a prestigious individual or a means of communication to many individuals in a group (one-to-many); e) from the elderly to the younger individuals in a population (many-to-one) [ 10 , 11 ]. Furthermore, according to Berkes [ 12 ], cultural traits can be interpreted as “ products of generations of intelligent reflection tested in the rigorous laboratory of survival, whose persistence is evidence of their power .” However, the prevalence of a cultural trait in a population can either remain stable or change dramatically over time, depending on the type of information, transmission mechanisms, population size and the environmental context [ 4 , 9 , 13 ]. These can be classified as cultural norms (e.g., continuous cultural traits), knowledge about resources, techniques and tools (e.g., quantitative cultural traits ) and cultural innovations (e.g., novel cultural traits) [ 6 , 14 , 15 ]. The body of knowledge about plants and treatments used for a therapeutic purpose constitutes a quantitative cultural trait [ 6 , 14 , 15 ]. If this trait were preserved in human adaptive memory, it could potentially enhance the population's quality of life [ 1 – 3 ]. In this context, the structure, function, type and frequency of trait transmission could have a vital role [ 1 , 6 , 16 – 18 ]. The collection of medicinal plants, related treatments, and knowledgeable users can be regarded as the primary components of a medical botanical system within a community [ 5 , 19 ]. Patterns related to the selection of medicinal plants in pharmacopoeias have shown a marked prevalence of certain botanical families. This convergence phenomena in usage are attributed to the action of some factors alone or acting in combination: i) the presence of prominent organoleptic characteristics [ 16 , 17 , 20 – 22 ], ii) being associated with a doctrine of signatures [ 23 ], iii) therapeutic effectiveness [ 24 , 25 ], iv) environmental availability and accessibility [ 26 ]. The organization of plant use based on these principles generates a tendency to form a specific group mostly used by the majority of users. From the perspective of evolutionary ethnobiology and cultural evolution, it can be associated with the “consensus within diversity theory” (see also [ 27 ]) and “cultural attraction theory (see, e.g., [ 4 ]) [ 5 ]. Moving to a more specific perspective, regarding medical botanical systems in particular, it is termed “core kit” [ 18 , 28 , 29 ], “ensemble” [ 30 , 31 ], or “plant complexes” [ 32 ]. Concerning the process of incorporating new plants, according to some theories, it is interpreted as an event that allows expanding the “stock” of therapeutic possibilities [ 18 ]. It has been proposed that the phenotype and organoleptic characteristics of frequently used plants (i.e., the core) would serve as prototypes for the incorporation of new plants into the medicinal system [ 18 , 32 – 34 ]. In summary, the integration of new plants would be incorporated by connecting them to the core, and their few connections with the rest of the system would place them in a peripheral location, resulting in less widespread transmission of knowledge [ 16 – 18 , 22 , 27 , 35 , 31 ]. Finally, this pattern of medicinal system growth is consistent with a core-periphery structure reported in other natural systems [ 36 – 40 ]. To describe and explain this phenomenon, we propose a methodological approach using network analysis. This approach is based on graph theory and provides a relevant framework for understanding complex systems in general [41, 42] and ethnobotanical knowledge systems in particular [35, 43, 44]. An advantage of the methodological approach is its ability to describe and explain network data through abstract models, enabling the representation of a complex reality [41, 42, 45] and visualizing how individuals engage with natural resources to identify patterns of interaction [35, 46]. Furthermore, this approach provides a valuable tool for ethnographic studies seeking to unravel this complexity [47]. In this way, network analyses enable s the detection of patterns in complex systems by representing a wide range of natural phenomena, such as interaction networks [41]. This approach has allowed us to reveal the presence of a core-periphery structure in a medical botanical system. This is composed of a basic set or “Kit” of medicinal plants that form a structural core [18, 27, 35, 31, 48]. Furthermore, this phenomenon of organizing interactions around a structural core is widely recognized in various complex systems [37, 38, 40, 49, 50]. In addition, this has allowed us to understand the determinants of agent cohesion and dispersion in an interaction network, as well as their effects on system functioning [49]. From a network theory and analytical perspective, the existence of a core corresponds to a group of nodes strongly interconnected with each other, which cannot be easily fragmented or divided, thus providing structural stability. The stability of the core also promotes the stability of peripheral nodes and, consequently, the overall system [37, 38, 40, 49, 50]. Medical botanical systems can be analyzed as graphs represented in a matrix where the rows contain the plants and the columns contain knowledgeable users [35, 43, 51, 52]. And particularly, medicinal plants can be related to the users as well as the diseases or therapeutic objectives associated with them [44]. This suggests, from the theory of complex networks, there are strong reasons to expect core-periphery assemblies in ethnobotany and it is noteworthy that ethnobotanical theory itself has predicted this pattern [18, 21, 31, 43]. Furthermore, the presence of a core structure is related to the system's stability and its ability to withstand disturbances, ensuring the population's health [ 38 ]. This property becomes particularly relevant when considering the erosion processes reported in traditional knowledge systems experienced by some populations [44, 53, 54]. Thus discovering a structure that provides stability could contribute to biocultural conservation goals [55]. Our hypothesis suggests that adaptive memory preserves quantitative cultural traits, such as those associated with a select group of beneficial medicinal plants present in medical botanical systems, which contribute to safeguard and improve population quality of life. Medicinal plants provide a structural core in these systems, and their higher frequency of use promotes their inclusion in this subgroup. These plants have been identified by most users, establishing a consensus core. By reaffirming the information that is circulating among users, the consensus in usage ensures that it is transmitted culturally and strengthens the system in the face of possible disturbance scenarios. From this hypothesis, we predict first: medicinal botanical systems present a “basic kit” of plants that is visualized as a structural core; second: core plants are more resistant and third: the versatility of uses is associated with belonging in the structural core, with prioritization of knowledge transmission, and therefore, with the conservation of their use. Finally, this study's goals are to provide quantitative support for a core-periphery structure within the medicinal botanical system in a locality in northern Uruguay and attempts to advance our understanding of the potential factors contributing to the formation of a core composed of frequently selected and used medicinal plants. Materials and methods Ethnobotanical Data Collection Snowball sampling was used to identify and interview forty-four participants, next to analyze the knowledge network concerning medicinal plants among the residents of “Parque Regional Quebradas del Norte” in Rivera, Uruguay (also see Castiñeira Latorre et al. for details on ethnographic methodology [44]). For all participants, we used semi-structured interviews and participant observation method to record local knowledge, thirty medicinal uses (etic categories) were documented, derived from 109 reported uses by the interlocutors (emic categories), and 159 species of medicinal plants were identified [42, 56–59]. Data analysis Based on this information, an affiliation matrix was compiled, consisting of interlocutors (rows of the matrix) and medicinal plants (columns of the matrix) (see also [44]). The matrix X is of dimension I x J, corresponding to the number of nodes. The element Xij represents the link that connects elements i and j. In the case of these two graphs, they do not have weighted entries. Therefore, if a link exists between i and j, the value of Xij is 1, and if there is no link, Xij is 0. These characteristics result in a bipartite network where connections only occur between nodes of different types [42]. A bipartite network was constructed consisting of interlocutors (n = 44) and medicinal plants (n = 159). For the network of interlocutors vs. medicinal plants, the presence of a core-periphery structure was determined, identifying the species corresponding to each component [ 40 ]. Using the “kcores” function from the “sna” package (Tools for Social Network Analysis), the largest of the k-cores based on degrees was identified. With the purpose of assessing the robustness of the network components, both the core and the periphery, and testing the hypothesis that the core of the medicinal system is preserved over time [ 18 ], we conducted a computational experiment. This experiment involved the random removal of interlocutors without replacement in the network of interlocutors vs. species of medicinal plants. After each removal, the proportion of plant species remaining within the core and periphery was quantified. Subsequently, the number of uses of medicinal plants corresponding to the core was compared to the uses of medicinal plants corresponding to the periphery. The means were compared using unpaired t-tests. Lastly, a linear regression was employed to determine if plants that have been allocated show more uses have also been assigned more interlocutors. The open-source R software system [60] was used for all studies. The “bipartite” package (which visualizes bipartite networks and calculates certain ecological indicators), the “sna” package (which offers tools for social network analysis), and the “igraph” package (which offers network analysis and visualization). Results A core-periphery structure was shown by the bipartite network, which included 44 interlocutors and 159 species of medicinal plants (Fig. 1 , Table 1 Supplementary material). Thirty-three of the 159 species that the interlocutors mentioned comprise the core structure (Fig. 1 , Table 1 Supplementary material). This result is consistent with the first prediction of the hypothesis. The result of the removal experiment is consistent with the second prediction of the hypothesis and shows that the proportion of species that remain in the system is always higher in the core than in the periphery of the network (Fig. 2 ). With the removal of two interlocutors, 52% of the species are maintained in the core and 12% in the periphery. In this sense, with only 10 interlocutors, 100% of the core species and 50% of the periphery species are maintained. Regarding the third prediction of the hypothesis, it is observed that the plants in the core have a higher cumulative number of uses mentioned by interlocutors, in comparison to the plants in the periphery. (X̅ core =20.88 vs. X̅ periphery =3.89; t [33, 12] = 7.9; p < 0.001; Fig. 3 ). When exploring the relationship between popularity and versatility, a positive linear relationship was detected between the number of assigned uses and the number of interlocutors (F [1, 157] = 1225; r2 = 0.89; p < 0.0001; Fig. 4 ). Discussion Through the analysis of social [46, 47, 58, 61–63] and biological [37, 40, 50, 64, 65] network structures, the relationship between different concepts formalized in network theory [41, 45, 66] was studied, and we found a correspondence in the context of ethnobotanical theory [18, 21, 35, 31, 67, 68]. Thus, with this conceptual framework and using network analysis tools, it was possible to identify a core-periphery structure in the network of the medical botanical system of the population at the “Parque Regional Quebradas del Norte”, Rivera, Uruguay. As it has been observed, most people agree to use a particular group of plants, within a wider range of medicinal plants, as therapeutic options for the treatment of various ailments and/or maintaining health. The “Consensus within Diversity” theory [ 27 ] has been summarized by Ferreira Júnior et al. [ 18 ] and is demonstrated in various ethnobotanical research (e.g., [ 21 , 31 , 35 ]. Moreover, a quantitative review of medicinal plants in the Patagonian Andes reports that in Mapuche medicine, there would be a regional pattern, a product of the local diversity of botanical species, and a global one shared with other pharmacopoeias, referring to the presence of a “cluster of consensual knowledge.” Continuing with the statement that “the Mapuche pharmacopoeia possesses a common body of botanical knowledge” [ 22 ]. In addition, these observations have also been highlighted for food botanical resources, such as the diversity of ethnospecies of “yuca” ( Manihot esculenta ) in rural communities [ 35 ]. Thus, employing network analysis methods, the reflections based on extensive ethnobotanical surveys agree with the core-periphery structure observed in this study [ 18 , 28 – 32 ]. Now, if the core-periphery structure is supported, the question arises: What might be the possible reasons for consensus on the core species? Suppose our adaptive memory is selective and has limitations in recalling the vast diversity of medicinal plants available in an environment [ 1 – 3 ]. In that case, we would likely remember particular plants or prototypes [ 16 ]. These prototype plants have characteristics that allow them to be recognized as medicinal, including organoleptic properties that would act as mnemonic resources, facilitating their memorization and guiding experimentation toward new, similar plants [ 16 , 17 ]. Additionally, these mnemonic resources help organize and anchor complex information, primarily in early learning (i.e., childhood and youth, [ 2 , 3 ]). Aroma and taste are powerful tools that induce the association of these prototype plants with the concepts of health and disease [ 21 , 22 ]. Within this study, the vast majority of plants in the core possess at least one of these organoleptic characteristics: aroma and bitter taste, which are associated with the concentration of essential oils and alkaloids, respectively [69–71]. However, other characteristics that medicinal plants possess could be relevant for the integration of these cores [ 18 ]. In this context, Bennett [ 23 ] highlights the significance of the mnemonic resource associated with the doctrine of signatures, in which individuals recognize the therapeutic value of plants based on specific morphological characteristics. Despite receiving numerous criticisms (see Bennett [ 23 ]), there are studies that demonstrate significant correlations between morphology and cultural attributes associated with use [ 21 ]. While the pharmacological efficacy of a plant may be an important factor remaining in the core, there could also be other attributes associated with effectiveness or cultural value [72, 73]. Hilgert [ 31 ] showed sociocultural illnesses associated with the breaking of taboos and supernatural elements, where medicinal plants act as a vehicle for the curative procedures practised by rural doctors and traditional healers. In summary, while the vast majority of plants within the core exhibit distinct ive organoleptic properties associated with pharmacological efficacy, potentially making them “the best models” for experimentation (sensu Leonti [ 17 ]). The loss of traditional knowledge is a phenomenon extensively documented in literature [45, 53, 67, 74]. Different causes are attributed to it, such as rural depopulation or the loss of qualified interlocutors (e.g. elders, shamans) [ 18 ]. In this work, we report to a medical botanical system that presents a core-periphery structure, where the core is resistant to the loss of plants (i.e. people knowledge) when we remove interlocutors (i.e. erosion or loss of traditional knowledge). Therefore, the basic kit of medicinal plants is preserved and resistant to disturbances in a medical botanical system with this structure (see, e.g., [ 18 ]). According to the preserved nature of medicinal knowledge within the core plants, it is expected that these plants are prioritized in the transmission of ethnobotanical knowledge through a combination of various mechanisms (e.g., vertical, horizontal, radial, and oblique) [17, 75, 76]. In addition, it is observed that there is a similar proportion of native and adventitious plants, which is expected in multicultural societies [77, 78]. As it has been observed, core knowledge is the most widespread and develops by a gradual accumulation of information through generations, providing the medical system resilience to knowledge loss [51, 53]. According to Ferreira Júnior et al. [ 18 ], peripheral plants have the ability to persist throughout the system, serving as a reservoir or "stock" of information. Periphery plants could act as therapeutic inventions in reaction to new illnesses [1, 9, 77, 78]. Our findings indicate that the plants which make up the core structure are the most well-known to the population (i.e., popular) and are attributed a higher number of treatments for various diseases (i.e., versatile plants). This allows us to interpret that versatility is an important characteristic for inclusion in the core structure. A similar pattern has been reported for communities in northeastern Brazil [79, 80]. This trend could be explained by the presence of various secondary metabolites in some plants, which could be related to their action on a variety of therapeutic targets [70]. Therefore, plants used for a specific treatment without adverse health effects are perceived by populations as safe plants, facilitating greater experimentation and use for a wider range of treatments over time [ 18 ]. Furthermore, a relationship has been found between the severity of diseases and their associated treatments [ 9 ], as well as the maintenance of this information in adaptive memory. These findings confirm results obtained from the core structure, where popular medicinal plant therapies are used to treat frequent less severe ailments [79] and are maintained in the disturbance scenarios. However, for a comprehensive interpretation of the phenomenon, future considerations should include aspects related to pharmacological efficacy and culturally perceived effectiveness [55, 72, 73, 81]. Finally, we explored a region with high priority for biocultural conservation (sensu Gavin et al. [81]) employing an interdisciplinary approach that integrated various fields of knowledge, including ethnographic, ecological and network analysis [43]. Our goal was to investigate the concept of “Consensus within Diversity” as proposed by Barrett [ 27 ] and further developed by Ferreira Júnior et al. [ 18 ]. The outcomes of this investigation provide valuable insights for developing strategies that prioritize efforts and underscore the urgency of preserving prototype plants within the core as well as potential innovations within the periphery of medicinal botanical systems. On one hand, core plants act as prototypes for other potential species, and their loss by extinction could significantly affect the system's resilience. On the other hand, peripheral plants are highly vulnerable but may offer possibilities for therapeutic innovations, especially in the face of changing scenarios. Addressing this dilemma could benefit from an examination of medicinal treatments associated with core and peripheral plants, taking into consideration disease prevalence within medicinal botanical systems. In the face of environmental change conservation efforts should be concentrated on vulnerable to extinction species mainly of the core and on people's knowledge from the peripheral plants as a biocultural conservation strategy (sensu Gavin et al. [81]). Declarations Acknowledgements: We are specially grateful to all the residents of the Bioma Pampa-Quebradas del Norte Biosphere Reserve, Rivera, Uruguay. ECL is particularly grateful for the theoretical and methodological contributions of the Matías Arim, Director of the Ecology Department at the Centro Universitario Regional del Este, CURE, Universidad de la República, and the María Leila Pochettino, Director of the Ethnobotany and Applied Botany Laboratory of the Facultad de Ciencias y Museo de la Universidad Nacional de La Plata, Argentina. Author contributions: EC designed the article and drafted the main text with the assistance of AB and AC. EC conducted field studies and various stages of research, obtaining results, including outreach activities in communities. AB and AC performed network analyses. All authors have read and approved the current manuscript version. Funding: This work was partially funded by the foreign postgraduate Scholarship Program of the Agencia Nacional de Investigación e Innovación (ANII), Uruguay. Availability of data and materials: All data used in this review can be found in the additional files. Ethics approval and consent to participate: Not applicable. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests. References Boyd R, Richerson P, Henrich J. The cultural niche: Why social learning is essential for human adaptation. PNAS. 2011;108:10918–25. Nairne JS, Pandeirada JNS. Adaptive Memory. Psych Science Perspectives. 2016; 11(4):496-511. Nairne JS, Pandeirada JNS. Adaptive memory: Is survival processing special? J Mem Lang. 2008;59(3):377–85. 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R Core Team. R: a language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing; 2016. Miceli J, Guerrero S. Redes libres de escala y su uso en el análisis de datos etnográficos: el caso de la comunidad Tehuelche del Chalía. 2005. Argentina Emperaire L, Olivera J. Redes sociales y diversidad agrícola en la Amazonía brasileña: un sistema multicéntrico. In: Pochettino ML, Ladio AH, Arenas P. Editors. Tradiciones y transformaciones en etnobotánica. Argentina: Cyted Prog;2010. 180–5. Pautasso M, Aistara G, Barnaud A, Caillon S, Clouvel P, Coomes OT, et al. Seed exchange networks for agrobiodiversity conservation. A review. Agron Sustain Dev. 2012;151–175. Borthagaray A, Arim M, Marquet P. Inferring species roles in metacommunity structure from species co-ocurrence networks. Proc R Soc. 2014. 281(1792):20141425 Borthagaray AI, Barreneche JM, Abades S, Arim M. Modularity along organism dispersal gradients challenges a prevailing view of abrupt transitions in animal landscape perception. Ecography. 2014;37:564–71. Marquitti FMD, Guimarães PR, Pires MM, Bittencourt LF. Modular: software for the autonomous computation of modularity in large network sets. Ecography. 2014;37(3):221–4. Reyes-García V, Guèze M, Luz AC, Paneque-Gálvez J, Macía MJ, Orta-Martínez M, et al. Evidence of traditional knowledge loss among a contemporary indigenous society. Evol Hum Behav. 2013;34(4):249–57. Diaz-Reviriego I, González-Segura L, Fernández-Llamazares A, Howard PL, Molina JL, Reyes-García V. Social organization influences the exchange and species richness of medicinal plants in Amazonian homegardens. Ecol Soc. 2015;21. Piastri M, Orfila L, Pardías P. Facultad de Química, Universidad de la República. 2017. In Tesauro de plantas medicinales. https://www.fq.edu.uy/?q=es/node/260. Accessed 20 Jan 2020. Alonso Paz E, Bassagoda MJ, Ferreira F. Yuyos: uso racional de las plantas medicinales. Fin de Siglo. Montevideo; 2008. 222 p. Santin JR, Lemos M, Júnior LCK, Niero R, de Andrade SF. Antiulcer effects of Achyrocline satureoides (Lam.) DC (Asteraceae) (Marcela), a folk medicine plant, in different experimental models. J Ethnopharmacol. 2010;130(2):334–9. de Medeiros PM, Ladio AH, Albuquerque UP. Local Criteria for Medicinal Plant Selection. 2015. In: Albuquerque UP, de Medeiros P, Casas A. (eds) Evolutionary Ethnobiology. Springer, Cham. https://doi.org/10.1007/978-3-319-19917-7_11 Menendez-Baceta G, Aceituno-Mata L, Reyes-García V, Tardío J, Salpeteur M, Pardo-de Santayana M. The importance of cultural factors in the distribution of medicinal plant knowledge: A case study in four Basque regions. J Ethnopharmacol. 2015;161:116–27. Albuquerque UP. Re-examining hypotheses concerning the use and knowledge of medicinal plants: a study in the Caatinga vegetation of NE Brazil. J Ethnobiol Ethnomed. 2006;2:30. Cavalli-Sforza LL, Feldman MW. Cultural transmission and evolution: a quantitative approach. 1981. Princeton, Princeton University Press. Hewlett BS, Cavalli-Sforza LL. Cultural transmission among Aka Pygmies. Am Anthropol. 1986; 88:922-934. Hanazaki N, Pieroni A, Ludwinsky RH, Gonçalves MC, Prakofjewa J, Peroni N, et al. People’s migrations and plants for food: a review for fostering sustainability. Sustain Earth Rev. 2023; 6(1):9. Vandebroek I, Balick MJ. Globalization and loss of plant knowledge: challenging the paradigm. PLoS One. 2012; 7(5):e37643. Santoro FR, Ferreira Júnior WS, Araújo TA de S, Ladio AH, Albuquerque UP. Does plant species richness guarantee the resilience of local medical systems? A perspective from utilitarian redundancy. PLoS One 2015; 10(3):e0119826. Lozano A, Araújo EL, Medeiros MFT, Albuquerque UP. The apparency hypothesis applied to a local pharmacopoeia in the Brazilian northeast. J Ethnobiol Ethnomed. 2014;10(1):2. Gavin MC, McCarter J, Mead A, Berkes F, Stepp JR, Peterson D, et al. Defining biocultural approaches to conservation. Trends Ecol Evol. 2015;30(3):140–5. Additional Declarations No competing interests reported. Supplementary Files Table1.docx Cite Share Download PDF Status: Published Journal Publication published 05 Nov, 2024 Read the published version in Journal of Ethnobiology and Ethnomedicine → Version 1 posted Editorial decision: Revision requested 14 Feb, 2024 Reviews received at journal 14 Feb, 2024 Reviewers agreed at journal 07 Feb, 2024 Reviews received at journal 06 Jan, 2024 Reviewers agreed at journal 30 Dec, 2023 Reviewers agreed at journal 28 Dec, 2023 Reviewers invited by journal 27 Dec, 2023 Editor assigned by journal 27 Dec, 2023 Submission checks completed at journal 27 Dec, 2023 First submitted to journal 22 Dec, 2023 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. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-3793784","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":264457125,"identity":"cc183417-53bc-4122-bb10-84519a3dc7ae","order_by":0,"name":"Elena Castiñeira Latorre","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2klEQVRIiWNgGAWjYDACdijNxt4AJA0siNDCDKEk2HgOgLRIkKCFQSIBQhME/MzMxx78qKmr45N8fnXDjwIJBv727gS8WiSb2dINe44dlmCTzim72QN0mMSZsxvwajE4zGMmzcB2AKQl7QYPUIuBRC5+LfaH+b9JM/yrk2CTPJN28w8xWgyYedikGduYJdgk2I/dJsoWicNsZpK9fYcl23hy2G7LGEjwEPQLf3vzM4kf3+r45duPP7v55o+NHH97L34tSIDHAEwSqxwE2B+QonoUjIJRMApGEAAAqLk8qfHfMqAAAAAASUVORK5CYII=","orcid":"","institution":"Universidad de la República","correspondingAuthor":true,"prefix":"","firstName":"Elena","middleName":"Castiñeira","lastName":"Latorre","suffix":""},{"id":264457126,"identity":"58378be0-94b4-4216-a853-36541560ac87","order_by":1,"name":"Ana Borthagaray","email":"","orcid":"","institution":"Universidad de la República","correspondingAuthor":false,"prefix":"","firstName":"Ana","middleName":"","lastName":"Borthagaray","suffix":""},{"id":264457127,"identity":"fa688732-51f7-4620-8913-98ffc6415146","order_by":2,"name":"Andrés Canavero","email":"","orcid":"","institution":"Universidad de la República","correspondingAuthor":false,"prefix":"","firstName":"Andrés","middleName":"","lastName":"Canavero","suffix":""}],"badges":[],"createdAt":"2023-12-22 20:29:07","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3793784/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3793784/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1186/s13002-024-00739-z","type":"published","date":"2024-11-05T15:57:02+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":49088846,"identity":"0fe663d7-bf31-44af-b7ab-8de5bcb39f87","added_by":"auto","created_at":"2024-01-03 01:28:38","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":765170,"visible":true,"origin":"","legend":"\u003cp\u003eThe Bioma Pampa-Quebradas del Norte Biosphere Reserve's ethnobotanical network, showing the species list that represents the core structure, links between interlocutors and medicinal plants, and a core-periphery structure.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-3793784/v1/9a3ac21458f5fe242819e403.png"},{"id":49088845,"identity":"ae277bf1-e602-4c92-b8b0-cf40d3f6516f","added_by":"auto","created_at":"2024-01-03 01:28:38","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":48700,"visible":true,"origin":"","legend":"\u003cp\u003eExperiments of removal. The percentage of core network species as a function of interlocutors count is shown by the dotted line, while the solid line shows the relationship with periphery species.\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-3793784/v1/eca3db643158d6bf7a60114d.png"},{"id":49088844,"identity":"11741348-f3d1-40d4-95a8-8f468eeb852f","added_by":"auto","created_at":"2024-01-03 01:28:38","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":43651,"visible":true,"origin":"","legend":"\u003cp\u003eBox plot shows the mean, standard deviations, and outliers of the number of uses for each species in the core and periphery of the interlocutors/medicinal plant network.\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-3793784/v1/1867f1d855860c9f92b94e09.png"},{"id":49089462,"identity":"c7702901-4d45-444a-a100-f2552fc0375e","added_by":"auto","created_at":"2024-01-03 01:36:43","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":75992,"visible":true,"origin":"","legend":"\u003cp\u003eCorrelation between the number of assigned uses and the number of interlocutors. Each point represents a medicinal species (n=159).\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-3793784/v1/545818ed2faa7d8e79bca454.png"},{"id":68749797,"identity":"ecfe437b-f01c-4cd6-85b9-eac64edce636","added_by":"auto","created_at":"2024-11-11 16:05:10","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1231521,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3793784/v1/e9709cb8-64b5-4d88-81a4-07ce12442126.pdf"},{"id":49088848,"identity":"f98035f3-ea6d-4070-97ea-efdf86bfe6ac","added_by":"auto","created_at":"2024-01-03 01:28:39","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":69847,"visible":true,"origin":"","legend":"","description":"","filename":"Table1.docx","url":"https://assets-eu.researchsquare.com/files/rs-3793784/v1/1daf048d50a4f71d8269afc9.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Core-periphery structure of a medicinal botanical system in Uruguay","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThroughout the course of adaptation, humans have developed cognitive processes to confront the difficulties presented by nature. As a result, natural selection has played a crucial role in shaping individuals' psychological structure, enabling them to remember experiences and enhance their ability to survive. Sensations and perceptions are naturally interpreted to create a response to difficult and stimulating environments [\u003cspan additionalcitationids=\"CR2 CR3\" citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]. Nairne \u0026amp; Pandeirada [\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e] propose that experiences are generationally remembered, integrated into an adaptive memory system. It has been documented that cultural features respond to situations of risk, partner preference, illness prevention, healing processes and illness therapies [\u003cspan additionalcitationids=\"CR6 CR7 CR8\" citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e]. These cultural traits are transmitted within a population through various pathways or mechanisms of knowledge diffusion: a) from parents to children (vertical); b) among peers of the same generation (horizontal); c) between generations, excluding the parenteral route (oblique); d) from a prestigious individual or a means of communication to many individuals in a group (one-to-many); e) from the elderly to the younger individuals in a population (many-to-one) [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]. Furthermore, according to Berkes [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e], cultural traits can be interpreted as \u0026ldquo;\u003cem\u003eproducts of generations of intelligent reflection tested in the rigorous laboratory of survival, whose persistence is evidence of their power\u003c/em\u003e.\u0026rdquo; However, the prevalence of a cultural trait in a population can either remain stable or change dramatically over time, depending on the type of information, transmission mechanisms, population size and the environmental context [\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]. These can be classified as cultural norms (e.g., continuous cultural traits), knowledge about resources, techniques and tools (e.g., quantitative cultural traits\u003cb\u003e)\u003c/b\u003e and cultural innovations (e.g., novel cultural traits) [\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe body of knowledge about plants and treatments used for a therapeutic purpose constitutes a quantitative cultural trait [\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e]. If this trait were preserved in human adaptive memory, it could potentially enhance the population's quality of life [\u003cspan additionalcitationids=\"CR2\" citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e]. In this context, the structure, function, type and frequency of trait transmission could have a vital role [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e, \u003cspan additionalcitationids=\"CR17\" citationid=\"CR36\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]. The collection of medicinal plants, related treatments, and knowledgeable users can be regarded as the primary components of a medical botanical system within a community [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e].\u003c/p\u003e \u003cp\u003ePatterns related to the selection of medicinal plants in pharmacopoeias have shown a marked prevalence of certain botanical families. This convergence phenomena in usage are attributed to the action of some factors alone or acting in combination: i) the presence of prominent organoleptic characteristics [\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e16\u003c/span\u003e, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e, \u003cspan additionalcitationids=\"CR21\" citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e], ii) being associated with a doctrine of signatures [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e], iii) therapeutic effectiveness [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e], iv) environmental availability and accessibility [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe organization of plant use based on these principles generates a tendency to form a specific group mostly used by the majority of users. From the perspective of evolutionary ethnobiology and cultural evolution, it can be associated with the \u0026ldquo;consensus within diversity theory\u0026rdquo; (see also [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]) and \u0026ldquo;cultural attraction theory (see, e.g., [\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]) [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]. Moving to a more specific perspective, regarding medical botanical systems in particular, it is termed \u0026ldquo;core kit\u0026rdquo; [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e], \u0026ldquo;ensemble\u0026rdquo; [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e], or \u0026ldquo;plant complexes\u0026rdquo; [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]. Concerning the process of incorporating new plants, according to some theories, it is interpreted as an event that allows expanding the \u0026ldquo;stock\u0026rdquo; of therapeutic possibilities [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]. It has been proposed that the phenotype and organoleptic characteristics of frequently used plants (i.e., the core) would serve as prototypes for the incorporation of new plants into the medicinal system [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan additionalcitationids=\"CR33\" citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e]. In summary, the integration of new plants would be incorporated by connecting them to the core, and their few connections with the rest of the system would place them in a peripheral location, resulting in less widespread transmission of knowledge [\u003cspan additionalcitationids=\"CR17\" citationid=\"CR36\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e]. Finally, this pattern of medicinal system growth is consistent with a core-periphery structure reported in other natural systems [\u003cspan additionalcitationids=\"CR37 CR38 CR39\" citationid=\"CR37\" class=\"CitationRef\"\u003e36\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e40\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eTo describe and explain this phenomenon, we propose a methodological approach using network analysis. This approach is based on graph theory and provides a relevant framework for understanding complex systems in general [41, 42] and ethnobotanical knowledge systems in particular [35, 43, 44]. An advantage of the methodological approach is its ability to describe and explain network data through abstract models, enabling the representation of a complex reality [41, 42, 45] and visualizing how individuals engage with natural resources to identify patterns of interaction [35, 46]. Furthermore, this approach provides a valuable tool for ethnographic studies seeking to unravel this complexity [47].\u003c/p\u003e \u003cp\u003eIn this way, network analyses enable\u003cb\u003es\u003c/b\u003e the detection of patterns in complex systems by representing a wide range of natural phenomena, such as interaction networks [41]. This approach has allowed us to reveal the presence of a core-periphery structure in a medical botanical system. This is composed of a basic set or \u0026ldquo;Kit\u0026rdquo; of medicinal plants that form a structural core [18, 27, 35, 31, 48]. Furthermore, this phenomenon of organizing interactions around a structural core is widely recognized in various complex systems [37, 38, 40, 49, 50]. In addition, this has allowed us to understand the determinants of agent cohesion and dispersion in an interaction network, as well as their effects on system functioning [49]. From a network theory and analytical perspective, the existence of a core corresponds to a group of nodes strongly interconnected with each other, which cannot be easily fragmented or divided, thus providing structural stability. The stability of the core also promotes the stability of peripheral nodes and, consequently, the overall system [37, 38, 40, 49, 50].\u003c/p\u003e \u003cp\u003eMedical botanical systems can be analyzed as graphs represented in a matrix where the rows contain the plants and the columns contain knowledgeable users [35, 43, 51, 52]. And particularly, medicinal plants can be related to the users as well as the diseases or therapeutic objectives associated with them [44]. This suggests, from the theory of complex networks, there are strong reasons to expect core-periphery assemblies in ethnobotany and it is noteworthy that ethnobotanical theory itself has predicted this pattern [18, 21, 31, 43]. Furthermore, the presence of a core structure is related to the system's stability and its ability to withstand disturbances, ensuring the population's health [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e38\u003c/span\u003e]. This property becomes particularly relevant when considering the erosion processes reported in traditional knowledge systems experienced by some populations [44, 53, 54]. Thus discovering a structure that provides stability could contribute to biocultural conservation goals [55].\u003c/p\u003e \u003cp\u003eOur hypothesis suggests that adaptive memory preserves quantitative cultural traits, such as those associated with a select group of beneficial medicinal plants present in medical botanical systems, which contribute to safeguard and improve population quality of life. Medicinal plants provide a structural core in these systems, and their higher frequency of use promotes their inclusion in this subgroup. These plants have been identified by most users, establishing a consensus core. By reaffirming the information that is circulating among users, the consensus in usage ensures that it is transmitted culturally and strengthens the system in the face of possible disturbance scenarios. From this hypothesis, we predict first: medicinal botanical systems present a \u0026ldquo;basic kit\u0026rdquo; of plants that is visualized as a structural core; second: core plants are more resistant and third: the versatility of uses is associated with belonging in the structural core, with prioritization of knowledge transmission, and therefore, with the conservation of their use. Finally, this study's goals are to provide quantitative support for a core-periphery structure within the medicinal botanical system in a locality in northern Uruguay and attempts to advance our understanding of the potential factors contributing to the formation of a core composed of frequently selected and used medicinal plants.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003ch2\u003eEthnobotanical Data Collection\u003c/h2\u003e \u003cp\u003eSnowball sampling was used to identify and interview forty-four participants, next to analyze the knowledge network concerning medicinal plants among the residents of \u0026ldquo;Parque Regional Quebradas del Norte\u0026rdquo; in Rivera, Uruguay (also see Casti\u0026ntilde;eira Latorre et al. for details on ethnographic methodology [44]). For all participants, we used semi-structured interviews and participant observation method to record local knowledge, thirty medicinal uses (etic categories) were documented, derived from 109 reported uses by the interlocutors (emic categories), and 159 species of medicinal plants were identified [42, 56\u0026ndash;59].\u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003eBased on this information, an affiliation matrix was compiled, consisting of interlocutors (rows of the matrix) and medicinal plants (columns of the matrix) (see also [44]). The matrix X is of dimension I x J, corresponding to the number of nodes. The element Xij represents the link that connects elements i and j. In the case of these two graphs, they do not have weighted entries. Therefore, if a link exists between i and j, the value of Xij is 1, and if there is no link, Xij is 0. These characteristics result in a bipartite network where connections only occur between nodes of different types [42]. A bipartite network was constructed consisting of interlocutors (n\u0026thinsp;=\u0026thinsp;44) and medicinal plants (n\u0026thinsp;=\u0026thinsp;159). For the network of interlocutors vs. medicinal plants, the presence of a core-periphery structure was determined, identifying the species corresponding to each component [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e40\u003c/span\u003e]. Using the \u0026ldquo;kcores\u0026rdquo; function from the \u0026ldquo;sna\u0026rdquo; package (Tools for Social Network Analysis), the largest of the k-cores based on degrees was identified. With the purpose of assessing the robustness of the network components, both the core and the periphery, and testing the hypothesis that the core of the medicinal system is preserved over time [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e], we conducted a computational experiment. This experiment involved the random removal of interlocutors without replacement in the network of interlocutors vs. species of medicinal plants. After each removal, the proportion of plant species remaining within the core and periphery was quantified. Subsequently, the number of uses of medicinal plants corresponding to the core was compared to the uses of medicinal plants corresponding to the periphery. The means were compared using unpaired t-tests. Lastly, a linear regression was employed to determine if plants that have been allocated show more uses have also been assigned more interlocutors. The open-source R software system [60] was used for all studies. The \u0026ldquo;bipartite\u0026rdquo; package (which visualizes bipartite networks and calculates certain ecological indicators), the \u0026ldquo;sna\u0026rdquo; package (which offers tools for social network analysis), and the \u0026ldquo;igraph\u0026rdquo; package (which offers network analysis and visualization).\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eA core-periphery structure was shown by the bipartite network, which included 44 interlocutors and 159 species of medicinal plants (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Table\u0026nbsp;1 Supplementary material). Thirty-three of the 159 species that the interlocutors mentioned comprise the core structure (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Table\u0026nbsp;1 Supplementary material). This result is consistent with the first prediction of the hypothesis. The result of the removal experiment is consistent with the second prediction of the hypothesis and shows that the proportion of species that remain in the system is always higher in the core than in the periphery of the network (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). With the removal of two interlocutors, 52% of the species are maintained in the core and 12% in the periphery. In this sense, with only 10 interlocutors, 100% of the core species and 50% of the periphery species are maintained. Regarding the third prediction of the hypothesis, it is observed that the plants in the core have a higher cumulative number of uses mentioned by interlocutors, in comparison to the plants in the periphery. (X̅\u003csub\u003ecore\u003c/sub\u003e=20.88 \u003cem\u003evs.\u003c/em\u003e X̅\u003csub\u003eperiphery\u003c/sub\u003e=3.89; t\u003csub\u003e[33, 12]\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;7.9; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). When exploring the relationship between popularity and versatility, a positive linear relationship was detected between the number of assigned uses and the number of interlocutors (F\u003csub\u003e[1, 157]\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;1225; r2\u0026thinsp;=\u0026thinsp;0.89; p\u0026thinsp;\u0026lt;\u0026thinsp;0.0001; Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThrough the analysis of social [46, 47, 58, 61\u0026ndash;63] and biological [37, 40, 50, 64, 65] network structures, the relationship between different concepts formalized in network theory [41, 45, 66] was studied, and we found a correspondence in the context of ethnobotanical theory [18, 21, 35, 31, 67, 68]. Thus, with this conceptual framework and using network analysis tools, it was possible to identify a core-periphery structure in the network of the medical botanical system of the population at the \u0026ldquo;Parque Regional Quebradas del Norte\u0026rdquo;, Rivera, Uruguay.\u003c/p\u003e\n\u003cp\u003eAs it has been observed, most people agree to use a particular group of plants, within a wider range of medicinal plants, as therapeutic options for the treatment of various ailments and/or maintaining health. The \u0026ldquo;Consensus within Diversity\u0026rdquo; theory [\u003cspan class=\"CitationRef\"\u003e27\u003c/span\u003e] has been summarized by Ferreira J\u0026uacute;nior et al. [\u003cspan class=\"CitationRef\"\u003e18\u003c/span\u003e] and is demonstrated in various ethnobotanical research (e.g., [\u003cspan class=\"CitationRef\"\u003e21\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e31\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e35\u003c/span\u003e]. Moreover, a quantitative review of medicinal plants in the Patagonian Andes reports that in Mapuche medicine, there would be a regional pattern, a product of the local diversity of botanical species, and a global one shared with other pharmacopoeias, referring to the presence of a \u0026ldquo;cluster of consensual knowledge.\u0026rdquo; Continuing with the statement that \u0026ldquo;the Mapuche pharmacopoeia possesses a common body of botanical knowledge\u0026rdquo; [\u003cspan class=\"CitationRef\"\u003e22\u003c/span\u003e]. In addition, these observations have also been highlighted for food botanical resources, such as the diversity of ethnospecies of \u0026ldquo;yuca\u0026rdquo; (\u003cem\u003eManihot esculenta\u003c/em\u003e) in rural communities [\u003cspan class=\"CitationRef\"\u003e35\u003c/span\u003e]. Thus, employing network analysis methods, the reflections based on extensive ethnobotanical surveys agree with the core-periphery structure observed in this study [\u003cspan class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e28\u003c/span\u003e\u0026ndash;\u003cspan class=\"CitationRef\"\u003e32\u003c/span\u003e]. Now, if the core-periphery structure is supported, the question arises: What might be the possible reasons for consensus on the core species? Suppose our adaptive memory is selective and has limitations in recalling the vast diversity of medicinal plants available in an environment [\u003cspan class=\"CitationRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan class=\"CitationRef\"\u003e3\u003c/span\u003e]. In that case, we would likely remember particular plants or prototypes [\u003cspan class=\"CitationRef\"\u003e16\u003c/span\u003e]. These prototype plants have characteristics that allow them to be recognized as medicinal, including organoleptic properties that would act as mnemonic resources, facilitating their memorization and guiding experimentation toward new, similar plants [\u003cspan class=\"CitationRef\"\u003e16\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e17\u003c/span\u003e]. Additionally, these mnemonic resources help organize and anchor complex information, primarily in early learning (i.e., childhood and youth, [\u003cspan class=\"CitationRef\"\u003e2\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e3\u003c/span\u003e]). Aroma and taste are powerful tools that induce the association of these prototype plants with the concepts of health and disease [\u003cspan class=\"CitationRef\"\u003e21\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e22\u003c/span\u003e]. Within this study, the vast majority of plants in the core possess at least one of these organoleptic characteristics: aroma and bitter taste, which are associated with the concentration of essential oils and alkaloids, respectively [69\u0026ndash;71]. However, other characteristics that medicinal plants possess could be relevant for the integration of these cores [\u003cspan class=\"CitationRef\"\u003e18\u003c/span\u003e]. In this context, Bennett [\u003cspan class=\"CitationRef\"\u003e23\u003c/span\u003e] highlights the significance of the mnemonic resource associated with the doctrine of signatures, in which individuals recognize the therapeutic value of plants based on specific morphological characteristics. Despite receiving numerous criticisms (see Bennett [\u003cspan class=\"CitationRef\"\u003e23\u003c/span\u003e]), there are studies that demonstrate significant correlations between morphology and cultural attributes associated with use [\u003cspan class=\"CitationRef\"\u003e21\u003c/span\u003e]. While the pharmacological efficacy of a plant may be an important factor remaining in the core, there could also be other attributes associated with effectiveness or cultural value [72, 73]. Hilgert [\u003cspan class=\"CitationRef\"\u003e31\u003c/span\u003e] showed sociocultural illnesses associated with the breaking of taboos and supernatural elements, where medicinal plants act as a vehicle for the curative procedures practised by rural doctors and traditional healers. In summary, while the vast majority of plants within the core exhibit distinct\u003cstrong\u003eive\u003c/strong\u003e organoleptic properties associated with pharmacological efficacy, potentially making them \u0026ldquo;the best models\u0026rdquo; for experimentation (sensu Leonti [\u003cspan class=\"CitationRef\"\u003e17\u003c/span\u003e]).\u003c/p\u003e\n\u003cp\u003eThe loss of traditional knowledge is a phenomenon extensively documented in literature [45, 53, 67, 74]. Different causes are attributed to it, such as rural depopulation or the loss of qualified interlocutors (e.g. elders, shamans) [\u003cspan class=\"CitationRef\"\u003e18\u003c/span\u003e]. In this work, we report to a medical botanical system that presents a core-periphery structure, where the core is resistant to the loss of plants (i.e. people knowledge) when we remove interlocutors (i.e. erosion or loss of traditional knowledge). Therefore, the basic kit of medicinal plants is preserved and resistant to disturbances in a medical botanical system with this structure (see, e.g., [\u003cspan class=\"CitationRef\"\u003e18\u003c/span\u003e]). According to the preserved nature of medicinal knowledge within the core plants, it is expected that these plants are prioritized in the transmission of ethnobotanical knowledge through a combination of various mechanisms (e.g., vertical, horizontal, radial, and oblique) [17, 75, 76]. In addition, it is observed that there is a similar proportion of native and adventitious plants, which is expected in multicultural societies [77, 78]. As it has been observed, core knowledge is the most widespread and develops by a gradual accumulation of information through generations, providing the medical system resilience to knowledge loss [51, 53]. According to Ferreira J\u0026uacute;nior et al. [\u003cspan class=\"CitationRef\"\u003e18\u003c/span\u003e], peripheral plants have the ability to persist throughout the system, serving as a reservoir or \u0026quot;stock\u0026quot; of information. Periphery plants could act as therapeutic inventions in reaction to new illnesses [1, 9, 77, 78].\u003c/p\u003e\n\u003cp\u003eOur findings indicate that the plants which make up the core structure are the most well-known to the population (i.e., popular) and are attributed a higher number of treatments for various diseases (i.e., versatile plants). This allows us to interpret that versatility is an important characteristic for inclusion in the core structure. A similar pattern has been reported for communities in northeastern Brazil [79, 80]. This trend could be explained by the presence of various secondary metabolites in some plants, which could be related to their action on a variety of therapeutic targets [70]. Therefore, plants used for a specific treatment without adverse health effects are perceived by populations as safe plants, facilitating greater experimentation and use for a wider range of treatments over time [\u003cspan class=\"CitationRef\"\u003e18\u003c/span\u003e]. Furthermore, a relationship has been found between the severity of diseases and their associated treatments [\u003cspan class=\"CitationRef\"\u003e9\u003c/span\u003e], as well as the maintenance of this information in adaptive memory. These findings confirm results obtained from the core structure, where popular medicinal plant therapies are used to treat frequent less severe ailments [79] and are maintained in the disturbance scenarios. However, for a comprehensive interpretation of the phenomenon, future considerations should include aspects related to pharmacological efficacy and culturally perceived effectiveness [55, 72, 73, 81].\u003c/p\u003e\n\u003cp\u003eFinally, we explored a region with high priority for biocultural conservation (sensu Gavin et al. [81]) employing an interdisciplinary approach that integrated various fields of knowledge, including ethnographic, ecological and network analysis [43]. Our goal was to investigate the concept of \u0026ldquo;Consensus within Diversity\u0026rdquo; as proposed by Barrett [\u003cspan class=\"CitationRef\"\u003e27\u003c/span\u003e] and further developed by Ferreira J\u0026uacute;nior et al. [\u003cspan class=\"CitationRef\"\u003e18\u003c/span\u003e]. The outcomes of this investigation provide valuable insights for developing strategies that prioritize efforts and underscore the urgency of preserving prototype plants within the core as well as potential innovations within the periphery of medicinal botanical systems. On one hand, core plants act as prototypes for other potential species, and their loss by extinction could significantly affect the system\u0026apos;s resilience. On the other hand, peripheral plants are highly vulnerable but may offer possibilities for therapeutic innovations, especially in the face of changing scenarios. Addressing this dilemma could benefit from an examination of medicinal treatments associated with core and peripheral plants, taking into consideration disease prevalence within medicinal botanical systems. In the face of environmental change conservation efforts should be concentrated on vulnerable to extinction species mainly of the core and on people\u0026apos;s knowledge from the peripheral plants as a biocultural conservation strategy (sensu Gavin et al. [81]).\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe are specially grateful to all the residents of the Bioma Pampa-Quebradas del Norte Biosphere Reserve, Rivera, Uruguay. ECL is particularly grateful for the theoretical and methodological contributions of the Mat\u0026iacute;as Arim, Director of the Ecology Department at the Centro Universitario Regional del Este, CURE, Universidad de la Rep\u0026uacute;blica, and the Mar\u0026iacute;a Leila Pochettino, Director of the Ethnobotany and Applied Botany Laboratory of the Facultad de Ciencias y Museo de la Universidad Nacional de La Plata, Argentina.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eEC designed the article and drafted the main text with the assistance of AB and AC. EC conducted field studies and various stages of research, obtaining results, including outreach activities in communities. AB and AC performed network analyses. All authors have read and approved the current manuscript version.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was partially funded by the foreign postgraduate Scholarship Program of the Agencia Nacional de Investigaci\u0026oacute;n e Innovaci\u0026oacute;n (ANII), Uruguay.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and materials:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll data used in this review can be found in the additional files.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eBoyd R, Richerson P, Henrich J. 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Criterios ambientales y organol\u0026eacute;pticos en los patrones de selecci\u0026oacute;n y uso de plantas medicinales en una comunidad Mapuche de la Patagonia semi\u0026aacute;rida Argentina. In:Pochettino ML, Ladio AH, Arenas PM, editors. Tradiciones y transformaciones en etnobot\u0026aacute;nica. Argentina: CYTED-Programa Iberoamericano Ciencia y Tecnolog\u0026iacute;a para el Desarrollo; 2009. p. 286\u0026ndash;302. \u003c/li\u003e\n\u003cli\u003eBorgatti SP, Everett MG. Models of core/periphery structures. Soc Networks. 1999;21:375 95. \u003c/li\u003e\n\u003cli\u003eMiele V, Ramos‐Jiliberto R, V\u0026aacute;zquez DP. Core\u0026ndash;periphery dynamics in a plant\u0026ndash;pollinator network. Journal of Animal Ecology. 2020;89(7):1670\u0026ndash;7. \u003c/li\u003e\n\u003cli\u003eHenrich J, Broesch J. On the nature of cultural transmission networks: evidence from Fijian villages for adaptive learning biases. 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Routledge. https://doi.org/10.4324/9781849776073\u003c/li\u003e\n\u003cli\u003eEstomba D, Ladio A, Lozada M. Medicinal wild plant knowledge and gathering patterns in a Mapuche community from North-western Patagonia. J Ethnopharmacol. 2006;103(1):109\u0026ndash;19. \u003c/li\u003e\n\u003cli\u003eNoy C. Sampling Knowledge: The Hermeneutics of Snowball Sampling in Qualitative Research. Int J Soc Res Methodol. 2008; 11(4):327\u0026ndash;44. \u003c/li\u003e\n\u003cli\u003eAlbuquerque UP. A little bit of Africa in Brazil: ethnobiology experiences in the field of Afro-Brazilian religions. J Ethnobiol Ethnomed 2014;10(1):12. \u003c/li\u003e\n\u003cli\u003eR Core Team. R: a language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing; 2016. \u003c/li\u003e\n\u003cli\u003eMiceli J, Guerrero S. Redes libres de escala y su uso en el an\u0026aacute;lisis de datos etnogr\u0026aacute;ficos: el caso de la comunidad Tehuelche del Chal\u0026iacute;a. 2005. 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Springer, Cham. https://doi.org/10.1007/978-3-319-19917-7_11\u003c/li\u003e\n\u003cli\u003eMenendez-Baceta G, Aceituno-Mata L, Reyes-Garc\u0026iacute;a V, Tard\u0026iacute;o J, Salpeteur M, Pardo-de Santayana M. The importance of cultural factors in the distribution of medicinal plant knowledge: A case study in four Basque regions. J Ethnopharmacol. 2015;161:116\u0026ndash;27. \u003c/li\u003e\n\u003cli\u003eAlbuquerque UP. Re-examining hypotheses concerning the use and knowledge of medicinal plants: a study in the Caatinga vegetation of NE Brazil. J Ethnobiol Ethnomed. 2006;2:30. \u003c/li\u003e\n\u003cli\u003eCavalli-Sforza LL, Feldman MW. Cultural transmission and evolution: a quantitative approach. 1981. Princeton, Princeton University Press.\u003c/li\u003e\n\u003cli\u003eHewlett BS, Cavalli-Sforza LL. Cultural transmission among Aka Pygmies. Am Anthropol. 1986; 88:922-934.\u003c/li\u003e\n\u003cli\u003eHanazaki N, Pieroni A, Ludwinsky RH, Gon\u0026ccedil;alves MC, Prakofjewa J, Peroni N, et al. People\u0026rsquo;s migrations and plants for food: a review for fostering sustainability. Sustain Earth Rev. 2023; 6(1):9. \u003c/li\u003e\n\u003cli\u003eVandebroek I, Balick MJ. Globalization and loss of plant knowledge: challenging the paradigm. PLoS One. 2012; 7(5):e37643. \u003c/li\u003e\n\u003cli\u003eSantoro FR, Ferreira J\u0026uacute;nior WS, Ara\u0026uacute;jo TA de S, Ladio AH, Albuquerque UP. Does plant species richness guarantee the resilience of local medical systems? A perspective from utilitarian redundancy. PLoS One 2015; 10(3):e0119826.\u003c/li\u003e\n\u003cli\u003eLozano A, Ara\u0026uacute;jo EL, Medeiros MFT, Albuquerque UP. The apparency hypothesis applied to a local pharmacopoeia in the Brazilian northeast. J Ethnobiol Ethnomed. 2014;10(1):2. \u003c/li\u003e\n\u003cli\u003eGavin MC, McCarter J, Mead A, Berkes F, Stepp JR, Peterson D, et al. Defining biocultural approaches to conservation. Trends Ecol Evol. 2015;30(3):140\u0026ndash;5. \u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"journal-of-ethnobiology-and-ethnomedicine","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jeet","sideBox":"Learn more about [Journal of Ethnobiology and Ethnomedicine](http://ethnobiomed.biomedcentral.com/)","snPcode":"13002","submissionUrl":"https://submission.nature.com/new-submission/13002/3","title":"Journal of Ethnobiology and Ethnomedicine","twitterHandle":"@ethnobiomed","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Adaptive memory, Core-periphery structure, Medical Botanical System, Medicinal plants, Network analysis, Resilience, Traditional knowledge","lastPublishedDoi":"10.21203/rs.3.rs-3793784/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3793784/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cstrong\u003eBackground: \u003c/strong\u003eHuman evolution has granted upon individual’s cognitive mechanisms necessary for remembering experiences, vital for both survival and reproduction. These experiences manifest into cultural traits, influencing human culture, particularly in healthcare and maintenance. Studies regarding medicinal plants and treatments are integral to the study of the medical botanical system. Pharmacopeias highlight the prevalence of specific species widely used, aligning with the “consensus within diversity theory” in evolutionary ethnobiology.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMethods:\u003c/strong\u003e This study applies network analysis on the botanical system of “Parque Regional Quebradas del Norte”, Rivera, Uruguay.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eResults:\u003c/strong\u003e Study results demonstrate a core-periphery structure, with a tightly interconnected core resistant to fragmentation, ensuring structural stability. This study showed the existence of peripheral nodes in the entire system, improving resilience against disturbances and increasing population's health.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConclusion: \u003c/strong\u003eThe core species, renowned for their versatility and multiple medicinal uses, treating less severe ailments effectively. Additionally, core plants serve as prototypes for innovations. Their extinction poses a threat to the system's resilience. Conversely, peripheral plants, though vulnerable, offer possibilities for therapeutic innovations. In the face of environmental change, conservation efforts should prioritize species that are vulnerable to extinction, particularly within the core. Simultaneously, preserving knowledge associated with peripheral plants presents a bicultural conservation strategy, ensuring the botanical system's robustness amongst evolving ecological conditions.\u003c/p\u003e","manuscriptTitle":"Core-periphery structure of a medicinal botanical system in Uruguay","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-01-03 01:28:34","doi":"10.21203/rs.3.rs-3793784/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2024-02-14T13:12:14+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-02-14T12:01:16+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"0c2b4889-57e0-421e-b3f1-d6fd7607b2ee","date":"2024-02-07T07:46:39+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-01-06T14:43:59+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"2b415224-2cca-4c8c-8562-3107fe0694ea","date":"2023-12-30T08:18:41+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"464cd034-3c93-4527-b976-d7f046a1df94","date":"2023-12-28T12:50:39+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2023-12-28T04:26:37+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2023-12-27T23:18:28+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2023-12-27T23:18:28+00:00","index":"","fulltext":""},{"type":"submitted","content":"Journal of Ethnobiology and Ethnomedicine","date":"2023-12-22T20:15:04+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"journal-of-ethnobiology-and-ethnomedicine","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jeet","sideBox":"Learn more about [Journal of Ethnobiology and Ethnomedicine](http://ethnobiomed.biomedcentral.com/)","snPcode":"13002","submissionUrl":"https://submission.nature.com/new-submission/13002/3","title":"Journal of Ethnobiology and Ethnomedicine","twitterHandle":"@ethnobiomed","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"63f5080a-3d5b-4f62-bde5-11ff629b62cc","owner":[],"postedDate":"January 3rd, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-11-11T15:59:09+00:00","versionOfRecord":{"articleIdentity":"rs-3793784","link":"https://doi.org/10.1186/s13002-024-00739-z","journal":{"identity":"journal-of-ethnobiology-and-ethnomedicine","isVorOnly":false,"title":"Journal of Ethnobiology and Ethnomedicine"},"publishedOn":"2024-11-05 15:57:02","publishedOnDateReadable":"November 5th, 2024"},"versionCreatedAt":"2024-01-03 01:28:34","video":"","vorDoi":"10.1186/s13002-024-00739-z","vorDoiUrl":"https://doi.org/10.1186/s13002-024-00739-z","workflowStages":[]},"version":"v1","identity":"rs-3793784","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3793784","identity":"rs-3793784","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}