In the target article, Lew-Levy and Amir argue persuasively that human children are important agents of cultural innovation and transmission and that their role in cultural evolution should be more fully appreciated. However, to understand the origins of human culture and processes of cultural evolution more broadly, it is critical to also consider other species. It is well established that culturally transmitted forms of behaviour are common in many nonhuman animal (hereafter animal) species (Whiten, Reference Whiten2021). Here, we argue that, as in humans, juveniles are likely to play a crucial – though largely overlooked – role in this process. To support this argument, we draw on three lines of evidence: (1) juveniles are often innovators in animal culture; (2) there are ample opportunities for cultural transmission within juvenile groups; and (3) recent work points to potential pathways for the transmission of social information from young to old in animal taxa.
As Lew-Levy and Amir point out, a standard model of cultural transmission is one in which older individuals transmit cultural information to younger individuals. This transmission pathway is undoubtedly important in animal societies and will often be adaptive as older, more experienced individuals are likely to possess valuable skills and knowledge. Indeed, there are many examples of cultural transmission in animal systems that fit this model (e.g., Dugatkin & Godin, Reference Dugatkin and Godin1993; Coelho et al., Reference Coelho, Falótico, Izar, Mannu, Resende, Siqueira and Ottoni2015; Arbon et al., Reference Arbon, Hahn, McIvor and Thornton2023). Yet, this model does not fit all observations. Take, for example, what can arguably be considered one of the first and certainly among the most famous examples of cultural transmission in animals: sweet potato washing in Japanese macaques. This tradition, in which macaques clean grit off provisioned sweet potatoes in the sea, presumably to make them more palatable, originated with a young female named Imo and spread among community members. Remarkably, this behaviour spread upward, from Imo and her peers to older community members, including Imo’s mother (Kawai, Reference Kawai1965).
Across a diverse range of species, immature individuals – like Imo – are highly exploratory and innovative, and so may serve as valuable originators of cultural information (Biro et al., Reference Biro, Inoue-Nakamura, Tonooka, Yamakoshi, Sousa and Matsuzawa2003; Thornton & Malapert, Reference Thornton and Malapert2009; Morand-Ferron et al., Reference Morand-Ferron, Cole, Rawles and Quinn2011; Benson-Amram et al., Reference Benson-Amram, Heinen, Gessner, Weldele and Holekamp2014; Coelho et al., Reference Coelho, Falótico, Izar, Mannu, Resende, Siqueira and Ottoni2015; Carter et al., Reference Carter, Forss, Page and Ratcliffe2018). One reason may lie in the old adage that necessity is the mother of invention: juveniles are typically low-ranking members of their groups and may, therefore, need to find creative ways to secure resources (Morand-Ferron et al., Reference Morand-Ferron, Cole, Rawles and Quinn2011). For instance, field experiments show that juvenile great tits and blue tits are particularly likely to solve novel food puzzles (Morand-Ferron et al., Reference Morand-Ferron, Cole, Rawles and Quinn2011) while in vampire bats, meerkats and chimpanzees, youngsters are most likely to explore novel objects, foraging tasks or food types (Biro et al., Reference Biro, Inoue-Nakamura, Tonooka, Yamakoshi, Sousa and Matsuzawa2003; Thornton & Malapert, Reference Thornton and Malapert2009; Carter et al., Reference Carter, Forss, Page and Ratcliffe2018). Young, naive individuals are also likely to be particularly reliant on social learning (Arbon et al., Reference Arbon, Boogert, Jordan and Thornton2025a).
If juveniles innovate, peer-to-peer social learning could, in principle, lead to the formation of juvenile cultures akin to those seen in human societies. Relatively little research has explored this explicitly, though there is evidence of social learning among juveniles in certain species such as lemon and Port Jackson sharks (Guttridge et al., Reference Guttridge, van Dijk, Stamhuis, Krause, Gruber and Brown2013; Pouca et al., Reference Pouca, Heinrich, Huveneers and Brown2020). Many animals form age-structured societies in which youngsters have ample opportunities to interact with and learn from each other. For instance, juvenile creches in many bird species and playful social interactions among juveniles in primates and other mammals are likely to provide ample opportunities for the transmission of foraging skills and the acquisition of social conventions (Marzluff & Balda, Reference Marzluff and Balda1992; Wanker, Bernate, & Franck, Reference Wanker, Bernate and Franck1996; Perry et al., Reference Perry, Baker, Fedigan, GrosLouis, Jack, MacKinnon, Manson, Panger, Pyle and Rose2003).
Might the cultural innovations of juvenile animals spread to affect the behaviour of adults, as Lew-Levy and Amir describe in humans? In some species, strong biases towards learning from older or more dominant individuals may prevent the transmission of cultural information from juveniles. For instance, some authors speculate that such social learning biases may be responsible for restricting the accumulation of cultural traditions in chimpanzee societies (Kendal et al., Reference Kendal, Hopper, Whiten, Brosnan, Lambeth, Schapiro and Hoppitt2015). As such, juveniles – despite their innovativeness – could represent something of a cultural dead-end. Yet, there are at least three possible pathways via which this dead-end could be avoided. First, juveniles may retain their learned behaviour as they grow up. In tufted capuchin monkeys, for example, juveniles showed particularly high rates of nut-cracking behaviour. Upon reaching adulthood, other group members then copied their behaviour, leading to the transmission of nut-cracking through the group (Coelho et al., Reference Coelho, Falótico, Izar, Mannu, Resende, Siqueira and Ottoni2015). Second, some animals have been shown to learn from successful individuals irrespective of their age (e.g. Seppänen et al., Reference Seppänen, Forsman, Mönkkönen, Krams and Salmi2011; Aplin et al., Reference Aplin, Farine, Morand-Ferron, Cockburn, Thornton and Sheldon2015). Finally, some species may flexibly alter their social information-use strategies. For example, jackdaws typically gather information from adults, but in a recent field experiment, adults learned to show social tolerance towards juveniles, using them as a source of information indicating the presence of food (Arbon et al., Reference Arbon, Truskanov, Stott, McIvor and Thornton2025b). Such flexibility may well allow some animals to capitalise on the innovativeness of youngsters, facilitating the spread of culture through groups.
Despite the relative dearth of research, multiple lines of evidence suggest that youngsters are likely to play a critical role in cultural transmission not only in humans, as argued by Lew-Levi and Amir, but also in many other species. Research on nonhuman animals, therefore, offers great potential, not least because of the feasibility of controlled experiments in the laboratory and in the field, which would be ethically and logistically difficult to conduct with humans. An explicit focus on the potential for peer-to-peer transmission among juveniles and from juveniles to adults should be an urgent research priority, not only for our fundamental understanding of cultural evolution, but also applied work harnessing social learning to enhance conservation (Arbon et al., Reference Arbon, Boogert, Jordan and Thornton2025a). The discovery that social learning operates as a “second inheritance system” (alongside genetic transmission) from parents to offspring has revolutionised biology and anthropology (Whiten, Reference Whiten2017). If this system allows information to spread up as well as down the generations, it may be even more powerful and versatile than we realised.
In the target article, Lew-Levy and Amir argue persuasively that human children are important agents of cultural innovation and transmission and that their role in cultural evolution should be more fully appreciated. However, to understand the origins of human culture and processes of cultural evolution more broadly, it is critical to also consider other species. It is well established that culturally transmitted forms of behaviour are common in many nonhuman animal (hereafter animal) species (Whiten, Reference Whiten2021). Here, we argue that, as in humans, juveniles are likely to play a crucial – though largely overlooked – role in this process. To support this argument, we draw on three lines of evidence: (1) juveniles are often innovators in animal culture; (2) there are ample opportunities for cultural transmission within juvenile groups; and (3) recent work points to potential pathways for the transmission of social information from young to old in animal taxa.
As Lew-Levy and Amir point out, a standard model of cultural transmission is one in which older individuals transmit cultural information to younger individuals. This transmission pathway is undoubtedly important in animal societies and will often be adaptive as older, more experienced individuals are likely to possess valuable skills and knowledge. Indeed, there are many examples of cultural transmission in animal systems that fit this model (e.g., Dugatkin & Godin, Reference Dugatkin and Godin1993; Coelho et al., Reference Coelho, Falótico, Izar, Mannu, Resende, Siqueira and Ottoni2015; Arbon et al., Reference Arbon, Hahn, McIvor and Thornton2023). Yet, this model does not fit all observations. Take, for example, what can arguably be considered one of the first and certainly among the most famous examples of cultural transmission in animals: sweet potato washing in Japanese macaques. This tradition, in which macaques clean grit off provisioned sweet potatoes in the sea, presumably to make them more palatable, originated with a young female named Imo and spread among community members. Remarkably, this behaviour spread upward, from Imo and her peers to older community members, including Imo’s mother (Kawai, Reference Kawai1965).
Across a diverse range of species, immature individuals – like Imo – are highly exploratory and innovative, and so may serve as valuable originators of cultural information (Biro et al., Reference Biro, Inoue-Nakamura, Tonooka, Yamakoshi, Sousa and Matsuzawa2003; Thornton & Malapert, Reference Thornton and Malapert2009; Morand-Ferron et al., Reference Morand-Ferron, Cole, Rawles and Quinn2011; Benson-Amram et al., Reference Benson-Amram, Heinen, Gessner, Weldele and Holekamp2014; Coelho et al., Reference Coelho, Falótico, Izar, Mannu, Resende, Siqueira and Ottoni2015; Carter et al., Reference Carter, Forss, Page and Ratcliffe2018). One reason may lie in the old adage that necessity is the mother of invention: juveniles are typically low-ranking members of their groups and may, therefore, need to find creative ways to secure resources (Morand-Ferron et al., Reference Morand-Ferron, Cole, Rawles and Quinn2011). For instance, field experiments show that juvenile great tits and blue tits are particularly likely to solve novel food puzzles (Morand-Ferron et al., Reference Morand-Ferron, Cole, Rawles and Quinn2011) while in vampire bats, meerkats and chimpanzees, youngsters are most likely to explore novel objects, foraging tasks or food types (Biro et al., Reference Biro, Inoue-Nakamura, Tonooka, Yamakoshi, Sousa and Matsuzawa2003; Thornton & Malapert, Reference Thornton and Malapert2009; Carter et al., Reference Carter, Forss, Page and Ratcliffe2018). Young, naive individuals are also likely to be particularly reliant on social learning (Arbon et al., Reference Arbon, Boogert, Jordan and Thornton2025a).
If juveniles innovate, peer-to-peer social learning could, in principle, lead to the formation of juvenile cultures akin to those seen in human societies. Relatively little research has explored this explicitly, though there is evidence of social learning among juveniles in certain species such as lemon and Port Jackson sharks (Guttridge et al., Reference Guttridge, van Dijk, Stamhuis, Krause, Gruber and Brown2013; Pouca et al., Reference Pouca, Heinrich, Huveneers and Brown2020). Many animals form age-structured societies in which youngsters have ample opportunities to interact with and learn from each other. For instance, juvenile creches in many bird species and playful social interactions among juveniles in primates and other mammals are likely to provide ample opportunities for the transmission of foraging skills and the acquisition of social conventions (Marzluff & Balda, Reference Marzluff and Balda1992; Wanker, Bernate, & Franck, Reference Wanker, Bernate and Franck1996; Perry et al., Reference Perry, Baker, Fedigan, GrosLouis, Jack, MacKinnon, Manson, Panger, Pyle and Rose2003).
Might the cultural innovations of juvenile animals spread to affect the behaviour of adults, as Lew-Levy and Amir describe in humans? In some species, strong biases towards learning from older or more dominant individuals may prevent the transmission of cultural information from juveniles. For instance, some authors speculate that such social learning biases may be responsible for restricting the accumulation of cultural traditions in chimpanzee societies (Kendal et al., Reference Kendal, Hopper, Whiten, Brosnan, Lambeth, Schapiro and Hoppitt2015). As such, juveniles – despite their innovativeness – could represent something of a cultural dead-end. Yet, there are at least three possible pathways via which this dead-end could be avoided. First, juveniles may retain their learned behaviour as they grow up. In tufted capuchin monkeys, for example, juveniles showed particularly high rates of nut-cracking behaviour. Upon reaching adulthood, other group members then copied their behaviour, leading to the transmission of nut-cracking through the group (Coelho et al., Reference Coelho, Falótico, Izar, Mannu, Resende, Siqueira and Ottoni2015). Second, some animals have been shown to learn from successful individuals irrespective of their age (e.g. Seppänen et al., Reference Seppänen, Forsman, Mönkkönen, Krams and Salmi2011; Aplin et al., Reference Aplin, Farine, Morand-Ferron, Cockburn, Thornton and Sheldon2015). Finally, some species may flexibly alter their social information-use strategies. For example, jackdaws typically gather information from adults, but in a recent field experiment, adults learned to show social tolerance towards juveniles, using them as a source of information indicating the presence of food (Arbon et al., Reference Arbon, Truskanov, Stott, McIvor and Thornton2025b). Such flexibility may well allow some animals to capitalise on the innovativeness of youngsters, facilitating the spread of culture through groups.
Despite the relative dearth of research, multiple lines of evidence suggest that youngsters are likely to play a critical role in cultural transmission not only in humans, as argued by Lew-Levi and Amir, but also in many other species. Research on nonhuman animals, therefore, offers great potential, not least because of the feasibility of controlled experiments in the laboratory and in the field, which would be ethically and logistically difficult to conduct with humans. An explicit focus on the potential for peer-to-peer transmission among juveniles and from juveniles to adults should be an urgent research priority, not only for our fundamental understanding of cultural evolution, but also applied work harnessing social learning to enhance conservation (Arbon et al., Reference Arbon, Boogert, Jordan and Thornton2025a). The discovery that social learning operates as a “second inheritance system” (alongside genetic transmission) from parents to offspring has revolutionised biology and anthropology (Whiten, Reference Whiten2017). If this system allows information to spread up as well as down the generations, it may be even more powerful and versatile than we realised.
Acknowledgements
We are grateful to our research groups – The Cooperation Lab at Boston College and the Wild Cognition Research Group at Exeter – for ongoing conversations that contributed to these ideas. We are especially grateful to Josh Arbon and Fred Ispanze for engaging discussions about animal culture. A.T. is supported by a Leverhulme Trust Research Grant (RGP-2020-170), and KM is supported by a Jacobs Foundation Research Fellowship.
Financial support
The authors have no funding do declare.
Competing interests
The authors declare no competing interest.