Champing refers to a conspicuous chewing or jaw-working motion performed in the absence of food and observed in social contexts in dogs and other canids. This short paper provides a descriptive ethological account of champing, interprets its function as a pacifying signal, and places it within established frameworks of social interaction and ontogenetic development. The behaviour is defined as a distinct ethological category based on the author’s long-term observations and comparative analysis.
Champing (also termed chomping) refers to a conspicuous, often audible chewing or jaw-working motion performed in the absence of food. In dogs and other canids, this behaviour is typically observed in social contexts. It is associated with affiliative intent, pacifying, insecurity, or submissiveness, depending on its intensity, timing, and accompanying signals.1
To the best of the author’s knowledge, this behaviour has not previously been described or formally defined as a distinct ethological category, despite being intermittently observed and subsumed under broader classes of pacifying or displacement behaviours.2
Across contexts, champing possesses a clear pacifying function. Pacifying behaviour (from Latin pacificare, pax = peace, facere = to make) comprises actions whose function is to reduce social tension, inhibit aggressive or dominant behaviour in another individual, or restore a state of social calm, as defined within an interactional framework of social behaviour (Hinde, 1976). In dogs, commonly described pacifying behaviours include licking, muzzle nudging, nose touching, pawing, yawning, body twisting, and head turning, all of which may be directed toward conspecifics or humans.
Champing is widely employed by canids in situations ranging from mild uncertainty to more pronounced social stress. Its acoustic and rhythmic properties appear to contribute to its communicative value, functioning as a low-risk, non-confrontational signal that advertises non-threatening intent (Lorenz, 1966).
Jane Goodall used to break a branch and pretend to chomp on it to pacify chimpanzees, showing some unease (photo by Derek Bryceson/National Geographic Creative).
Ontogenetically, champing has a plausible developmental basis. One of the earliest repetitive oral sounds in mammalian neonates is produced during suckling and is closely linked to satisfaction, warmth, and social contact. In puppies, early oral motor patterns tied to nursing occur in a context of comfort and need fulfilment. As development continues, elements of this behaviour are redirected into social functions, where champing helps turn uncomfortable or ambiguous interactions into more benign ones. Initially, the behaviour is closely tied to hunger reduction; later, it becomes separate from feeding and acquires a distinct communicative function (Hinde, 1982).
In adult dogs, champing is a clear and effective signal of affiliative or conciliatory intent. Similar patterns appear across mammals, where oral behaviours linked to nursing and sucking are associated with reduced arousal and resting states. This suggests early sensory–motor associations may keep a tension-reducing function throughout life.3
Comparable observations exist in primates. Jane Goodall reported deliberately mimicking chewing movements—such as breaking a twig and pretending to chew it—to pacify chimpanzees displaying signs of unease (Goodall, 1971).
In applied animal contexts, the author has often used champing with apparent success when interacting with dogs or horses, consistent with its proposed pacifying function.
Footnotes
In ethology, the formal identification and naming of behavioural patterns commonly precede their experimental isolation or quantification. Descriptive classification based on repeated observation, functional context, and comparative consistency has historically been a primary means by which distinct behavioural units are recognised, refined, and later subjected to experimental analysis. ↩︎
The present account is based on the author’s long-term ethological observations and comparative analyses of canine social behaviour, first described in Dog Language (Abrantes, 1986 and 1997). It is descriptive and functional in scope and does not claim experimental isolation, quantitative prevalence estimates, or phylogenetic exclusivity for champing behaviour. In the absence of prior formal treatment of this behaviour as a distinct category, these observations constitute the primary empirical basis for the description and interpretation presented here. ↩︎
Evidence for the calming or arousal-reducing effects of suckling and related oral behaviours in mammals is well established in the developmental and comparative literature. Studies of non-nutritive sucking and nursing behaviour report associations with reduced behavioural arousal and increased resting or quiet states in a range of species (e.g. Blass, 1980; Veissier et al., 2002). While these works do not address champing or later social signalling directly, they provide developmental support for the inference that early oral sensory–motor patterns may retain residual tension-reducing properties when redeployed in other behavioural contexts. ↩︎
References
Abrantes, R. (1997). Dog language: An encyclopedia of canine behaviour. Wakan Tanka, Publishers. (Original work published as Hundesprog in 1986).
Hinde, R. A. (1982). Ethology: Its nature and relations with other sciences. Oxford: Oxford University Press.
Lorenz, K. (1966). On aggression. London: Methuen.
Veissier, I., de Passillé, A. M., Després, G., Rushen, J., Charpentier, I., Ramirez de la Fe, A. R., & Pradel, P. (2002). Does nutritive and non-nutritive sucking reduce other oral behaviors and stimulate rest in calves? Journal of Animal Science, 80(10), 2574–2587. https://doi.org/10.1093/ansci/80.10.2574
Featured image: Champing behaviour has a pacifying function—attempting to turn an unpleasant situation into a pleasant one.
This article is originally written on April 12, 2017 and slightly edited on January 2, 2026.
Bonding is a central feature of social life in many animal species, yet the terms bonding, attachment, and imprinting are often conflated in both scientific and popular discourse. This article examines bonding as a biological and behavioral process, distinguishing its proximate mechanisms from its ultimate evolutionary functions. Focusing on two companion species with contrasting evolutionary ecologies—domestic dogs (Canis lupus familiaris) and domestic horses (Equus ferus caballus)—we compare how imprinting, attachment, and broader bonding processes emerge across development and social contexts. Drawing on ethology, comparative studies, and neurobiological research, we show that while early attachment relationships are developmentally constrained and species-specific, enduring social bonds are more flexibly shaped by shared experience, social regulation, and cooperative activity. We further argue that bonding should be understood not merely as an affiliative state, but as a regulatory process that supports coordination, stress regulation, and cooperation. By integrating evolutionary, developmental, and mechanistic perspectives, this comparative analysis clarifies key conceptual distinctions, common principles, and meaningful differences in social bonding across two different species.
Figure 1. Shared exposure to demanding and potentially stressful situations can strengthen social bonds through coordination, trust, and reciprocal regulation of behavior and arousal during joint activity. Search-and-rescue handlers and their dogs exemplify such cooperative relationships, which are shaped by repeated joint problem-solving under challenging conditions. Photo: Désirée Mallè, Alpine Rescue Team, and her dog.
Bonding—a Definition
In animal behavior, bonding refers to a biologically grounded process by which individuals—of the same or different species—develop stable, selective social relationships that are maintained over time. The primary adaptive functions of bonding include promoting coordination, cooperation, and mutual tolerance, thereby enhancing individual fitness and, in many cases, inclusive fitness.12
Bonding is expressed through recurrent interaction patterns that regulate access, proximity, and coordinated activity among partners. Its strength, duration, and symmetry vary widely across species and social contexts, ranging from transient affiliations to enduring, lifelong bonds.
Parent–Offspring Bonding and Attachment
The term attachment is used cautiously in ethology.3 When employed, it represents a functional and descriptive label for a particular regulatory organization of social behavior, rather than as a reference to inferred mental states or subjective experience. Ethological usage has historically emphasized parent–offspring relationships, especially during periods of functional dependency (Hinde, 1982; Bateson, 1994), and has cautioned against unqualified extension of the term to adult social relationships (Hinde, 1976; Silk, 2007).
In the present paper, we treat attachment as a specialized form of bonding, defined ethologically as a pattern of selective proximity regulation and context-dependent separation responses that serves regulatory and adaptive functions. While attachment is often most clearly expressed in filial contexts, it is not defined by age or developmental stage, but by its functional structure.
The most fundamental and extensively studied form of bonding occurs between parents and offspring. In this context, bonding frequently takes the form of attachment. Filial attachment promotes proximity maintenance, contributes to the regulation of distress during separation, and provides a secure base from which juveniles can explore their environment.
Filial attachment is typically most pronounced during periods of dependency and becomes less central as the juvenile attains functional independence. Nevertheless, early attachment-related interactions can exert enduring effects on later social behavior, stress responsiveness, and affiliative tendencies (Bowlby, 1982; Carter, 1998).45
In domestic dogs (Canis lupus familiaris), a well-documented sensitive period for social attachment occurs approximately between the third and tenth weeks of age. During this phase, puppies readily form selective social relationships with conspecifics and humans. Individuals deprived of typical social contact beyond roughly 14 weeks of age often show persistent alterations in social behavior, including reduced affiliative responsiveness and atypical interaction patterns relative to species- and population-specific norms (Scott & Fuller, 1965; Freedman et al., 1961).
Pair Bonding and Reproductive Cooperation
In many social species, males and females form pair bonds during courtship and mating. These bonds support coordinated reproductive behavior, including shared parental investment, mate guarding, or cooperative resource defense, thereby increasing the likelihood that shared genetic material is successfully transmitted to subsequent generations.
Pair bonding is functionally and evolutionarily favored in species where ecological conditions—e.g., prolonged offspring dependency, biparental care need, mate guarding, or dispersed resources—make sustained cooperation between reproductive partners more likely to increase the survival and reproductive success of their shared offspring, thereby enhancing the direct fitness of both parents (Clutton-Brock, 1991).
Pair bonds may incorporate attachment-like regulatory features, such as selective proximity and partner-specific buffering of stress. Yet, they are functionally distinct from filial attachment in that they primarily serve reproductive coordination rather than developmental dependency (Clutton-Brock, 1991).
Social Bonding Beyond Attachment
Among group-living animals, bonding also arises through repeated interaction, cohabitation, and shared ecological challenges. Such bonds need not involve attachment in the strict sense—that is, they may lack pronounced separation responses or stress-buffering functions—yet they remain stable and functionally significant.
Behaviors such as grooming, play, coalitionary support, and reciprocal food sharing are widespread mechanisms for maintaining social bonds. Shared intense experiences (Fig. 2), including coordinated responses to threats, are particularly effective in strengthening affiliative ties among adults, as they reduce uncertainty regarding partners’ reliability in critical contexts (Silk, 2007).6
Bonding should therefore not be understood solely as an affiliative or affective state, but also as a regulatory process emerging from shared coping with challenge and uncertainty, in which moderate, manageable stress can facilitate learning, coordination, and social cohesion (Carter, 1998; Insel & Young, 2001; Abrantes, 2025).
Figure 2. Cooperative interaction among members of Micromys minutus (photo by Cuttestpaw). Shared, demanding interactions can contribute to the formation and reinforcement of social bonds. Bonding, attachment, and imprinting represent distinct biological processes with different developmental timing and functions (see Table 1).
Neurobiological Substrates of Bonding and Attachment
At a proximate level, affiliative interactions are associated with neuroendocrine processes, notably the release of oxytocin, which modulates defensive responses and facilitates social approach, tolerance, and coordination (Carter, 1998; Insel & Young, 2001). These mechanisms support both broad social bonding and the more specific dynamics of attachment.
Attachment, however, also recruits systems involved in stress regulation and separation responses, including the hypothalamic–pituitary–adrenal (HPA) axis, endogenous opioid systems, and associated neural circuits that mediate distress and recovery during separation and reunion. These systems support the regulatory functions of attachment by modulating arousal, persistence, and recovery in the absence of social contact, distinguishing attachment from broader forms of social bonding (Carter, 1998; Insel & Young, 2001).
Figure 3. Conceptual relationships between bonding, imprinting, and attachment. Bonding represents the broad class of enduring affiliative social relationships that support cooperation, tolerance, and social regulation. Attachment constitutes a functionally specialized subset of bonding, characterized by selective proximity seeking and separation-related regulatory processes, most commonly expressed during periods of dependency but not restricted to them. Imprinting is a phase-sensitive learning process that can establish stable social preferences and thereby contribute to bond formation. The overlap between imprinting and attachment (imprinting-based attachment) reflects cases in which early learning supports the emergence of attachment relationships. The diagram emphasizes that bonding encompasses a broader range of social relationships than either imprinting or attachment alone, and that neither imprinting nor attachment is necessary for bonding to occur.
Bonding, Attachment, and Imprinting
Bonding is often discussed alongside imprinting, but the concepts are not interchangeable. While imprinting produces a bond, not all bonding involves imprinting, and not all bonds involve attachment7 (see Table 1).
Imprinting refers to a form of phase-sensitive learning that occurs during a restricted developmental window, is rapid, and appears largely independent of the immediate consequences of behavior. Some species are predisposed to acquire specific information—such as caregiver identity or species recognition—during these sensitive periods. This learning reflects evolved developmental programs rather than associative conditioning (Lorenz, 1935; Bateson, 1979).8
Attachment, by contrast, develops through ongoing interaction and experience. Although it often emerges during sensitive periods, it remains modifiable and is regulated by feedback from the caregiver–offspring relationship.
At the level of bonding as a general social process, fitness benefits may accrue through both direct and inclusive pathways, depending on whether bonds involve reproductive partners, kin, or non-kin; by contrast, pair bonding specifically enhances direct fitness via increased offspring survival.
Table 1—Terminological Comparison: Bonding, Attachment, and Imprinting
Term
Core Definition
Developmental Timing
Learning Mechanism
Typical Duration
Functional Role
Key References
Bonding
A biologically grounded process through which individuals form stable, selective social relationships maintained over time
Promotes coordination, cooperation, tolerance, and social stability; enhances individual and inclusive fitness
Carter, 1998; Insel & Young, 2001; Silk, 2007
Attachment
A functionally specialized form of bonding characterized by selective proximity regulation and context-dependent separation responses
Not developmentally restricted; often most pronounced during periods of dependency
Experience-dependent learning supporting proximity regulation, stress modulation, and partner-specific responses
Typically long-lasting; expression may change across contexts and life stages
Regulates proximity, buffers stress, and supports adaptive performance under vulnerability
Bowlby, 1969/1982; Carter, 1998; Insel & Young, 2001
Imprinting
A developmentally constrained learning process through which specific stimuli or social partners acquire enduring salience
Restricted sensitive or critical period
Rapid, often non-associative or weakly associative learning; relatively resistant to extinction
Typically long-lasting or irreversible
Biases later recognition, preference, or social orientation; may shape but does not constitute bonding or attachment
Lorenz, 1935; Bateson, 1979; Horn, 2004
Bonding and Attachment in Domestic Dogs
In domestic environments, dogs develop social bonds and, in many cases, attachment relationships through everyday interaction. Grooming, resting in proximity, play, coordinated vocal responses, and joint reactions to environmental disturbances contribute to the formation and maintenance of affiliative bonds.
Dogs form attachment relationships not only with conspecifics but also with humans, as evidenced by selective proximity regulation, stress modulation, and differential behavioral responses to familiar versus unfamiliar individuals (Topál et al., 1998; Gácsi et al., 2013). They may also form stable bonds with individuals of other species, such as household cats, reflecting the flexibility of canine social bonding systems.
Beyond early attachment formation, domestic dogs establish stable, selective social bonds with both conspecifics and humans. Preferred social partners, asymmetries in play solicitation, and selective proximity patterns cannot be explained by familiarity alone (Bradshaw & Nott, 1995; Cafazzo et al., 2010). These bonds are associated with measurable stress-buffering effects: the presence of a familiar human or canine partner reduces behavioral indicators of distress and attenuates physiological stress responses in challenging situations (Gácsi et al., 2013; Nagasawa et al., 2015).
Bond strength is not maintained by passive affiliation alone. Coordinated activity under mild challenge, including problem-solving and shared task engagement, appears particularly effective in reinforcing dog–human bonds, consistent with the view that shared regulation under manageable stress promotes durable social bonding (Gácsi et al., 2013; Nagasawa et al., 2015; Abrantes, 2025).
At a proximate level, affiliative interactions between dogs and humans—including mutual gaze, physical contact, and coordinated activity—are associated with increased oxytocin in both partners, supporting a conserved neuroendocrine substrate for social bonding across species (Odendaal & Meintjes, 2003; Nagasawa et al., 2015). While dogs readily form attachment relationships with humans, these attachments remain experience-dependent, shaped by consistency, predictability, and shared activity rather than by imprinting alone, reinforcing the distinction between early phase-sensitive learning and later-developing attachment bonds.
Bonding and Attachment in Domestic Horses
Domestic horses (Equus ferus caballus) are highly social, herd-living mammals in which bonding plays a central role in survival and welfare. In both free-ranging and managed populations, horses form stable affiliative relationships, expressed through preferred spatial proximity, synchronized activity, and allogrooming—a behavior closely associated with social tolerance and group cohesion (Waring, 2003; Budiansky, 1997). These bonds support collective vigilance and coordinated responses to potential threats, consistent with the horse’s evolutionary history as a socially obligate prey species.
The most prominent attachment relationship in horses is the mare–foal bond, which develops rapidly after birth and is essential for protection, learning, and early social development. Foals show selective following and behavioral disruption upon separation, while mares provide regulation through proximity and intervention. This attachment is strongest during early dependency and gradually diminishes as juveniles integrate into the wider social group. Evidence indicates that early social deprivation or premature separation can produce long-term effects on social behavior and responses to novelty and handling, highlighting the developmental importance of early attachment in horses (Søndergaard & Jago, 2010).
Beyond early development, adult horses form selective social bonds within the herd. Although these relationships do not necessarily meet strict attachment criteria—such as selective proximity regulation under acute stress—they are persistent and functionally significant. Preferred partners are associated with reduced behavioral indicators of fear and improved coping in challenging situations, suggesting that adult social bonding in horses serves a regulatory and stress-buffering function (Christensen et al., 2008; Lansade et al., 2008). Horses may also form bonds with humans; however, these relationships are best understood as experience-dependent and context-specific, shaped by predictability and shared activity rather than by imprinting or caregiver-style attachment (Waring, 2003; Budiansky, 1997).
In horses, as in dogs, bonding is more reliably reinforced through shared activity and coordinated responses to environmental challenges than through passive contact alone, consistent with a regulatory—rather than purely affiliative—interpretation of social bonding (Christensen et al., 2008; Lansade et al., 2008; Abrantes, 2025).
Comparative Perspective: Dogs and Horses
Dogs and horses illustrate how bonding and attachment processes are shaped by species-specific ecology while relying on shared biological principles (see Table 2). Dogs, as socially flexible carnivores shaped by intensive human-directed selection, readily form attachment relationships with humans that functionally resemble caregiver–offspring systems in key regulatory respects. Horses, as socially obligate prey animals, emphasize herd cohesion, mutual tolerance, and collective regulation, with attachment largely confined to early development and selected interspecific contexts.
In both species, enduring bonds are more reliably strengthened through shared experience and coordinated activity than through passive contact alone. These contrasts underscore the importance of distinguishing ultimate evolutionary function from proximate mechanisms, while demonstrating that bonding remains a general, cross-species process grounded in cooperation, regulation, and survival.
Table 2—Imprinting, Attachment, and Bonding in Domestic Dogs and Horses
Dimension
Dogs (Canis familiaris)
Horses (Equus ferus caballus)
Evolutionary niche
Social carnivore (predator) (Clutton-Brock, 1991)
Social herbivore (prey) (Waring, 2003)
Primary social ecology
Flexible social grouping; high social plasticity (Bradshaw & Nott, 1995)
Stable herd structure; social conservatism (Budiansky, 1997; Waring, 2003)
Imprinting
Clear sensitive period for social orientation (≈3–10 weeks), extendable to humans (Scott & Fuller, 1965; Freedman et al., 1961)
Primarily mare–foal recognition; limited beyond neonatal period (Waring, 2003)
Function of imprinting
Establishes early social orientation toward conspecifics and humans (Scott & Fuller, 1965)
Ensures early maternal recognition and cohesion (Waring, 2003)
Attachment (juvenile)
Strong puppy–caregiver attachment; selective proximity regulation and distress modulation (Topál et al., 1998)
Strong mare–foal attachment; declines with social integration (Søndergaard & Jago, 2010)
Attachment (adult)
Common toward humans; selective proximity regulation and stress buffering toward familiar partners (Topál et al., 1998; Gácsi et al., 2013)
Rare and context-specific; not typically expressed as proximity regulation under stress (Waring, 2003; Budiansky, 1997)
Bonding (conspecifics)
Selective social bonds; play and tolerance asymmetries (Bradshaw & Nott, 1995; Cafazzo et al., 2010)
Selective affiliative bonds; proximity and allogrooming (Waring, 2003)
Bonding (interspecific)
Stable, enduring bonds with humans common (Topál et al., 1998)
Bonds with humans experience-dependent and task-related (Budiansky, 1997)
Stress modulation by social partners
Strong; familiar humans or dogs reduce behavioral and physiological stress (Gácsi et al., 2013; Odendaal & Meintjes, 2003)
Moderate; preferred partners reduce fear responses and improve coping (Christensen et al., 2008; Lansade et al., 2008)
Bonding and stress regulation
Shared exposure to manageable challenges strengthens bonds via mutual regulation (Gácsi et al., 2013; Abrantes, 2025)
Shared coping and coordinated activity strengthen bonds via stress buffering (Christensen et al., 2008; Abrantes, 2025)
Neuroendocrine correlates
Oxytocin associated with human–dog bonding and stress modulation (Odendaal & Meintjes, 2003; Nagasawa et al., 2015)
Less directly studied; stress modulation inferred behaviorally and physiologically (Lansade et al., 2008)
Role of shared activity
Central to bond strengthening (Bradshaw & Nott, 1995; Abrantes, 2025)
Central to bond strengthening (Budiansky, 1997; Abrantes, 2025)
Risk of anthropomorphic misinterpretation
High if attachment inferred beyond demonstrated regulatory criteria (Topál et al., 1998)
High if attachment inferred without evidence of proximity regulation under stress (Waring, 2003)
Note. References listed in each cell are representative primary or synthetic sources supporting the stated patterns, not an exhaustive review. The table contrasts dominant tendencies shaped by species-specific ecology (predator vs. prey) and domestication history; individual variation and contextual effects are expected in both species.
Practical Implications for Human–Animal Interaction
The distinctions developed in this paper—between bonding, imprinting, and attachment—have direct implications for how humans interact with companion animals in everyday contexts. First, recognizing that imprinting is developmentally constrained, while attachment and bonding are not, underscores the importance of early social experience, particularly in dogs and in the mare–foal relationship in horses. In dogs, early social deprivation during sensitive periods has been shown to produce long-lasting deficits in social behavior and adaptability (Freedman et al., 1961; Scott & Fuller, 1965), while in horses, early handling and the quality of the mare–foal relationship significantly influence later responses to humans and novel situations (Søndergaard & Jago, 2010). These findings indicate that missed or impoverished early social exposure cannot be fully compensated for later by affiliative contact alone.
Second, understanding bonding as an experience-dependent and regulatory process shifts the emphasis from passive affiliative gestures to shared activity. While behaviors such as petting or eye contact can support short-term social engagement, empirical work in dogs shows that attachment-related stress buffering and proximity regulation are more robustly expressed in contexts involving coordinated interaction and human participation (Topál et al., 1998; Gácsi et al., 2013). Similarly, studies in horses indicate that social buffering effects are most evident when animals face challenges in the presence of a familiar partner, rather than through proximity alone (Christensen et al., 2008; Lansade et al., 2008).
Third, the role of manageable stress and challenge in bonding suggests that optimal interaction does not require eliminating all difficulty. Moderate stress, when predictably regulated and socially mediated, can facilitate learning and social cohesion rather than undermine it (Carter, 1998; Insel & Young, 2001). This interpretation is consistent with comparative evidence showing that shared coping with environmental or task-related challenges strengthens affiliative relationships in both dogs and horses (Gácsi et al., 2013; Christensen et al., 2008), and aligns with a regulatory rather than hedonic understanding of bonding (Abrantes, 2025).
Finally, distinguishing bonding from attachment helps prevent anthropomorphic expectations. Dogs readily form attachment relationships with human partners that meet established behavioral criteria, including selective proximity regulation and stress buffering (Topál et al., 1998; Gácsi et al., 2013), whereas horses typically do not exhibit attachment patterns that map onto caregiver–offspring models, despite forming stable and meaningful social bonds (Waring, 2003; Budiansky, 1997). Recognizing these species-specific differences allows humans to interact more effectively and more respectfully with each animal, aligning expectations with biological and ecological realities rather than with human social norms (Clutton-Brock, 1991).
Conclusion
This analysis has aimed to clarify the concept of bonding by situating it within a comparative and evolutionary framework, while carefully distinguishing it from attachment and imprinting. Using domestic dogs and horses as case studies, we have shown that bonding is neither reducible to early phase-sensitive learning nor synonymous with attachment relationships, even when these processes overlap in development and function (Bateson, 1979; Bowlby, 1982). Rather, bonding emerges as a flexible, experience-dependent process grounded in repeated interaction, shared activity, and social regulation (Carter, 1998; Insel & Young, 2001).
The comparison between dogs and horses illustrates how species-specific ecology and domestication history shape the expression of social relationships. Dogs, as socially plastic carnivores selected for close cooperation with humans, readily form attachment relationships with human partners that persist into adulthood and meet established regulatory criteria (Topál et al., 1998; Gácsi et al., 2013). Horses, by contrast, as socially obligate prey animals, emphasize herd cohesion and selective affiliative bonds, with attachment most clearly expressed in early developmental contexts and selected interspecific situations (Waring, 2003; Budiansky, 1997). Despite these differences, both species demonstrate that enduring bonds are strengthened more by coordinated action and shared coping with challenge than by passive affiliation (Christensen et al., 2008; Gácsi et al., 2013).
More broadly, distinguishing ultimate evolutionary explanations from proximate bonding mechanisms helps avoid both anthropomorphism and unwarranted generalization (Hamilton, 1964; Tinbergen, 1963). Bonding can be favored by natural selection in species where it promotes cooperation, tolerance, survival, and reproductive success. Yet, it is instantiated through learning, social experience, and physiological regulation rather than through intention or moral sentiment. Recognizing this multi-level structure allows for a more precise and biologically grounded understanding of social relationships in companion animals. It provides a model that can be extended—cautiously and explicitly—to other social species.
Footnotes
Inclusive fitness refers to the total genetic contribution an individual makes to subsequent generations, including both direct reproduction and effects on the reproductive success of genetically related individuals, weighted by degree of relatedness. This concept explains how social behaviors that appear altruistic at the individual level can be favored by natural selection when they enhance the transmission of shared genes (Hamilton, 1964). ↩︎
Inclusive fitness provides an ultimate, evolutionary explanation for why bonding can be favored by natural selection; the formation and maintenance of bonds themselves depend on proximate mechanisms, including development, learning, social experience, and neuroendocrine regulation. ↩︎
Psychological frameworks of attachment, including the Strange Situation paradigm (Ainsworth et al., 1978) and its application to dogs by Topál et al. (1998), are cited here solely for their operational separation–reunion criteria. These approaches originate in human developmental psychology and have generated ongoing discussion regarding their scope and interpretation when applied across species or beyond early developmental contexts (Hinde, 1982; Wynne, 2004; Buller, 2005). In the present paper, their use is restricted to clearly defined behavioral patterns, without theoretical commitments concerning mental states or emotional experience. ↩︎
Bowlby explicitly characterizes attachment in biological terms: “Attachment behaviour is regarded as a class of social behaviour of an importance equivalent to that of mating behaviour and parental behaviour. It is held to have a biological function specific to itself […]” (Bowlby, 1982, p. 223). This formulation treats attachment as an evolved behavioral system defined by function rather than by species-specific expression. ↩︎
Carter summarizes the functional role of social attachment as follows: “[…] social attachments function to facilitate reproduction, provide a sense of security and reduce feelings of stress or anxiety” (Carter, 1998, p. 779). ↩︎
As a general evolutionary principle, Silk defines the conditions under which sociality evolves as follows: “[…] sociality evolves when the net benefits of close association with conspecifics exceed the costs” (Silk, 2007, p. 539). This formulation provides the ultimate-level framework within which affiliative behaviors such as grooming, play, and cooperative defense can be understood as mechanisms that increase the reliability and benefits of social partners. ↩︎
The statement that imprinting produces a bond refers to the fact that imprinting establishes a stable social preference or orientation toward a particular individual, class of individuals, or stimulus, thereby generating an affiliative relation. However, imprinting is only one possible developmental pathway to bonding. Many bonds—such as adult affiliative relationships, cooperative partnerships, or interspecific social bonds—arise through repeated interaction, shared experience, and social regulation outside any restricted sensitive period. Conversely, not all bonds involve attachment in the strict sense defined by selective proximity seeking and distress regulation under separation. Attachment represents a specific subset of bonds, typically associated with dependency and security regulation, whereas bonding is the broader category encompassing a range of affiliative and cooperative social relationships (Bowlby, 1982; Bateson, 1979; Carter, 1998). ↩︎
The term imprinting (original German Prägung) was introduced by Konrad Lorenz to describe a distinctive form of early learning observed in birds, characterized by rapid acquisition, restricted to a sensitive developmental period, and relatively independent of reinforcement (Lorenz, 1935). Early formulations emphasized the apparent irreversibility of imprinting effects; however, subsequent research has shown that while imprinting outcomes are often highly stable, they are not invariably permanent and may be modifiable under certain conditions, particularly with later experience or altered social environments (Bateson, 1979; Horn, 2004). Significantly, this qualification does not undermine the core concept. Instead, it reflects a broader shift away from rigid dichotomies between innate and learned behavior toward a developmental perspective in which evolved predispositions interact with experience. Although the term imprinting has declined in frequency relative to broader constructs such as early learning or developmental plasticity, it remains scientifically relevant as a label for phase-sensitive learning processes that are rapid, time-constrained, and shaped by species-specific developmental programs rather than by associative conditioning alone. ↩︎
Ainsworth, M. D. S., Blehar, M. C., Waters, E., & Wall, S. (1978). Patterns of attachment: A psychological study of the strange situation. Lawrence Erlbaum Associates. ISBN 9780470267079
Bowlby, J. (1982). Attachment and loss: Vol. 1. Attachment (2nd ed.). Basic Books. (Original work published 1969). ISBN 9780465005437
Bradshaw, J. W. S., & Nott, H. M. R. (1995). Social and communication behaviour of companion dogs. In J. Serpell (Ed.), The domestic dog: Its evolution, behaviour and interactions with people (pp. 115–130). Cambridge University Press. ISBN 9780521415293
Budiansky, S. (1997). The nature of horses: Exploring equine evolution, intelligence, and behavior. Free Press. ISBN 9780684827681
Buller, D. J. (2005). Adapting minds: Evolutionary psychology and the persistent quest for human nature. MIT Press. ISBN 9780262524608
Cafazzo, S., Valsecchi, P., Bonanni, R., & Natoli, E. (2010). Dominance in relation to age, sex, and competitive contexts in a group of free-ranging domestic dogs. Behavioral Ecology, 21(3), 443–455. https://doi.org/10.1093/beheco/arq001
Christensen, J. W., Malmkvist, J., Nielsen, B. L., & Keeling, L. J. (2008). Effects of a calm companion on fear reactions in naïve test horses. Equine Veterinary Journal, 40(1), 46–50. https://doi.org/10.2746/042516408X245171
Clutton-Brock, T. H. (1991). The evolution of parental care. Princeton University Press. ISBN 9780691024685
Gácsi, M., Maros, K., Sernkvist, S., Faragó, T., & Miklósi, Á. (2013). Human analogue safe haven effect of the owner: Behavioural and heart rate response to stressful social stimuli in dogs. PLoS ONE, 8(3), e58475. https://doi.org/10.1371/journal.pone.0058475
Horn, G. (2004). Pathways of the past: The imprint of memory. Nature Reviews Neuroscience, 5(2), 108–120. https://doi.org/10.1038/nrn1324
Insel, T. R., & Young, L. J. (2001). The neurobiology of attachment. Nature Reviews Neuroscience, 2(2), 129–136. https://doi.org/10.1038/35053579
Lansade, L., Bouissou, M.-F., & Erhard, H. W. (2008). Fearfulness in horses: A temperament trait stable across time and situations. Applied Animal Behaviour Science, 109(2–4), 355–373. https://doi.org/10.1016/j.applanim.2008.06.011
Nagasawa, M., Mitsui, S., En, S., et al. (2015). Oxytocin-gaze positive loop and the coevolution of human–dog bonds. Science, 348(6232), 333–336. https://doi.org/10.1126/science.1261022
Odendaal, J. S. J., & Meintjes, R. A. (2003). Neurophysiological correlates of affiliative behaviour between humans and dogs. The Veterinary Journal, 165(3), 296–301. https://doi.org/10.1016/S1090-0233(02)00237-X
Scott, J. P., & Fuller, J. L. (1965). Genetics and the social behavior of the dog. University of Chicago Press. ISBN 9780226743430
Silk, J. B. (2007). The adaptive value of sociality in mammalian groups. Philosophical Transactions of the Royal Society B: Biological Sciences, 362(1480), 539–559. https://doi.org/10.1098/rstb.2006.1994
Søndergaard, E., & Jago, J. G. (2010). The effect of early handling of foals on their reaction to handling, humans and novelty, and the foal–mare relationship.Applied Animal Behaviour Science, 123(3–4), 93–100. https://doi.org/10.1016/j.applanim.2010.01.006
Topál, J., Miklósi, Á., Csányi, V., & Dóka, A. (1998). Attachment behavior in dogs (Canis familiaris): A new application of Ainsworth’s Strange Situation Test. Journal of Comparative Psychology, 112(3), 219–229. https://doi.org/10.1037/0735-7036.112.3.219
Waring, G. H. (2003). Horse behavior (2nd ed.). William Andrew Publishing. ISBN 9780815514871
Wynne, C. D. L. (2004). Do animals think? Princeton University Press. ISBN 9780691118650
Conflict of Interest Statement The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Stress is often portrayed as harmful, yet moderate, acute stress can enhance learning, memory retention, and social bonding. Recent epigenetic research reveals that stress hormones modulate gene expression in key brain regions, strengthening memory consolidation and attentional processes. Unpleasant or intense experiences tend to form long-lasting memories, an adaptive mechanism for survival. Beyond cognition, stress can facilitate social bonding through oxytocin-mediated social buffering, as demonstrated in mammals, including domesticated dogs, although effects are highly context-dependent. Excessive or chronic stress, however, disrupts these processes, impairing memory, social interactions, and overall well-being. This paper emphasizes the nuanced, dual role of stress, highlighting its adaptive functions and underscoring the importance of understanding stress within an evolutionary and behavioral framework, not least because such understanding can inform more efficient animal behavior modification.
Stress Helps Learning and Bonding
A tough nut to crack is an everlasting memory that binds the parties together, and there is a reason for that. Moderate stress heightens arousal and sharpens attention, facilitating learning and the formation of durable memories (Roozendaal, McEwen, & Chattarji, 2009; McGaugh, 2015). Studies show that stress-related hormones and neuromodulators can also strengthen certain social bonds, depending on context, species, and prior history (Carter, 2014; Hostinar, Sullivan, & Gunnar, 2014).
Fig. 1 — Illustration of the hypothalamic-pituitary-adrenal (HPA) axis during the stress response: the hypothalamus detects stress and releases corticotropin-releasing hormone (CRH), which stimulates the pituitary gland to release adrenocorticotropic hormone (ACTH). ACTH triggers the adrenal glands to produce cortisol, the body’s key stress hormone. Cortisol’s effects on the body feed back to regulate this system, maintaining balance through a negative feedback loop.
The Term Stress Is Dangerously Ambiguous
We need to be careful, though. The term stress is dangerously ambiguous. Richard Shweder once described stress in a 1997 New York Times, Week in Review essay, as “a word that is as useful as a Visa card and as satisfying as a Coke. It’s non-committal and also non-committable.” Here, we adopt a biological definition:
Stress is the organism’s coordinated physiological response to a real or perceived challenge to homeostasis, involving the activation of the sympathetic nervous system and the hypothalamic–pituitary–adrenal axis to restore equilibrium (see fig. 1).
This distinction—between colloquial and biological uses—is crucial because the physiological and behavioral mechanisms engaged differ depending on whether the stressor is acute or chronic, controllable or uncontrollable. In this context, Koolhaas et al. (2011, p. 1291) propose that “the term ‘stress’ should be restricted to conditions where an environmental demand exceeds the natural regulatory capacity of an organism, in particular situations that include unpredictability and uncontrollability,” emphasizing the adaptive and context-dependent nature of the stress response (McEwen & Wingfield, 2010; Koolhaas et al., 2011).
What Is the Function of Stress?
Being an evolutionary biologist, when contemplating a mechanism, I always ask: “What is the function of that? What is that good for?” A mechanism can originate by chance (most do), but unless it provides the individual with some extra benefits in survival and reproduction, it will not spread in the population. From an evolutionary perspective, the stress response and the modulation of memory under stress increase the probability of survival (Nesse & Ellsworth, 2009; McEwen, Nasca, & Gray, 2016).
Why Do Unpleasant Memories Persist?
Emotionally intense, threatening, or highly arousing situations produce stronger, more persistent memory traces. Biologically, remembering potentially harmful events helps self-preservation. Negative or threatening events recruit the amygdala–hippocampal network more strongly, with the amygdala modulating hippocampal consolidation via noradrenergic and glucocorticoid-dependent mechanisms (Johansen, Cain, Ostroff, & LeDoux, 2011; McGaugh, 2015; LeDoux & Pine, 2016).
Fig. 2 — Sequence of events from exposure to a stressor through activation of the body’s physiological and behavioral stress response system (including the HPA axis), resulting in molecular and epigenetic changes such as DNA methylation and altered gene expression in stress-related genes. These epigenetic modifications influence future stress responsiveness and can affect health outcomes over the long term.
Epigenetic Effects
One of the most exciting scientific discoveries of late is the role of epigenetics (see fig. 2). Epigenetics—the study of modifications in gene activity that occur without altering the DNA sequence—has become central to contemporary models of learning and memory. Bird defines an epigenetic event as “the structural adaptation of chromosomal regions so as to register, signal or perpetuate altered activity states” (Bird, 2007, p. 398). Within this framework, attention focuses on activity-dependent chromatin modifications that occur during an individual’s lifetime rather than on transgenerational inheritance (Allis & Jenuwein, 2016). Mechanisms such as DNA methylation, histone acetylation, and related chromatin adjustments fine-tune gene expression in response to salient experiences, enabling the formation and stabilization of memory (Sweatt, 2013). Stress hormones act on mineralocorticoid and glucocorticoid receptors in hippocampal and amygdalar circuits, where they modulate plasticity and enhance the consolidation of significant events (Roozendaal, McEwen, & Chattarji, 2009; McEwen et al., 2012). Through interactions with noradrenergic projections from the locus coeruleus, glucocorticoids further shape these epigenetic regulators, influencing transcriptional programs essential for synaptic plasticity (Zovkic, Guzman-Karlsson, & Sweatt, 2013; Gray, Rubin, Hunter, & McEwen, 2014). These coordinated molecular processes, under moderate stress, enhance learning and contribute to the durability of highly arousing or threatening experiences.
Not All Stress Boosts Learning
Not all stress is productive for learning. Excessive stress produces the opposite effect. There is a difference between being stressed and stressed out. When stress becomes excessive or prolonged, the organism enters a state where immediate survival takes priority over other functions, and memory formation decreases. Chronic stress, in particular, undermines learning and cognitive function by disrupting hippocampal structure and impairing synaptic plasticity (de Kloet, Joëls, & Holsboer, 2005). These maladaptive effects highlight that stress is beneficial only within a moderate and context-dependent range; beyond that, it impairs both cognition and emotional regulation.
Stress and Bonding—A Delicate Balance
Stress does more than enhance memory; under certain conditions, it actively promotes social bonding. Oxytocin, a neuropeptide closely linked to affiliation, mediates this effect by dampening the HPA axis response during shared or moderate stress, thereby encouraging proximity and affiliative behaviors (Crockford, Deschner, & Wittig, 2017). In rodents, moderate stress enhances social-seeking behavior among cagemates via oxytocin signaling, though excessively threatening contexts abolish this effect (Burkett et al., 2015). Findings in rodents provide a foundation for understanding oxytocin-mediated bonding, which can also be observed in humans and domesticated dogs, albeit with species-specific nuances.
In domesticated dogs, exogenous oxytocin increases sociability toward humans and conspecifics, and social interactions raise endogenous oxytocin levels (Nagasawa et al., 2015). Just as humans bond emotionally through mutual gaze—a process mediated by oxytocin—Nagasawa et al. demonstrate that a similar gaze-mediated bonding exists between humans and dogs: “These findings support the existence of an interspecies oxytocin-mediated positive loop facilitated and modulated by gazing, which may have supported the coevolution of human-dog bonding by engaging common modes of communicating social attachment” (Nagasawa et al., 2015, p. 333). Longitudinal observations further show that chronic stress markers, such as hair cortisol, can synchronize between dogs and their owners, suggesting a deep physiological linkage (Sundman et al., 2020). Importantly, these bonding effects are highly context-dependent: moderate, predictable stress tends to facilitate affiliation, whereas excessive or prolonged stress may inhibit social bonding.
Caveats: Despite the fascinating discoveries mentioned above, we must be prudent in our conclusions. The effects of stress on bonding are highly context-dependent. Elevated cortisol in dogs can reflect excitement rather than distress (Nagasawa et al., 2015), and the benefits observed in rodents require non-threatening environments (Burkett et al., 2015). Oxytocin’s influence varies with social familiarity; stress may not enhance affiliation with strangers or weakly bonded partners (Crockford et al., 2017). Correlational studies, such as cortisol synchronization in dog–owner dyads, cannot prove causality, though they suggest physiological coupling that may support bonding under shared stress.
Conclusion
We need a balanced view of stress. Acute, manageable challenges—those that elicit adaptive stress responses—support attentional sharpening, facilitate memory consolidation, strengthen social bonds, and promote effective learning. These benefits are highly context-dependent: stress can enhance cognition and affiliation when moderate and predictable, but excessive or prolonged stress can overwhelm these systems, impairing memory, social interactions, and overall well-being. From an evolutionary perspective, stress serves a dual adaptive function—preparing individuals to respond to threats while reinforcing social bonds that increase survival odds. A nuanced understanding is therefore essential for interpreting behavior and guiding sound practice.
For animal trainers, these insights translate into a few practical guidelines. Animals benefit from gradual exposure to manageable, stress-eliciting challenges that promote resilience and adaptive coping. Training sessions should be calibrated so that the stress elicited remains within a range that facilitates attention and learning—enough to trigger mild HPA-axis activation, but not so intense as to be counter-productive. Moreover, designing training sessions that employ an appropriate level of stress can strengthen the trainer–animal bond by allowing the trainer to serve as a social buffer during mildly stressful tasks.
Featured picture: A tough nut to crack is an everlasting memory that binds the parties together (photo by unknown).
References
Allis, C. D., & Jenuwein, T. (2016). The molecular hallmarks of epigenetic control. Nature Reviews Genetics, 17(8), 487–500. https://doi.org/10.1038/nrg.2016.59
Burkett, J. P., Andari, E., Johnson, Z. V., Curry, D. C., de Waal, F. B. M., & Young, L. J. (2016). Oxytocin‑dependent consolation behavior in rodents. Science, 351(6271), 375–378. https://doi.org/10.1126/science.aac4785
Crockford, C., Deschner, T., & Wittig, R. M. (2017). The role of oxytocin in social buffering of stress: What do primate studies add? Current Topics in Behavioral Neurosciences, 30, 1–33. https://doi.org/10.1007/7854_2017_12
de Kloet, E. R., Joëls, M., & Holsboer, F. (2005). Stress and the brain: From adaptation to disease. Nature Reviews Neuroscience, 6(6), 463–475. https://doi.org/10.1038/nrn1683
Gray, J. D., Rubin, T. G., Hunter, R. G., & McEwen, B. S. (2014). Hippocampal gene expression changes underlying stress sensitization and recovery. Molecular Psychiatry, 19(11), 1171–1178. https://doi.org/10.1038/mp.2013.175
Hostinar, C. E., Sullivan, R. M., & Gunnar, M. R. (2014). Psychobiological mechanisms underlying the social buffering of the stress response: A review of animal models and human studies across development. Psychological Bulletin, 140(1), 256–282. https://doi.org/10.1037/a0032671
Johansen, J. P., Cain, C. K., Ostroff, L. E., & LeDoux, J. E. (2011). Molecular mechanisms of fear learning and memory. Cell, 147(3), 509–524. https://doi.org/10.1016/j.cell.2011.10.009
Koolhaas, J. M., Bartolomucci, A., Buwalda, B., de Boer, S. F., Flügge, G., Korte, S. M., … Fuchs, E. (2011). Stress revisited: A critical evaluation of the stress concept. Neuroscience & Biobehavioral Reviews, 35(5), 1291–1301. https://doi.org/10.1016/j.neubiorev.2011.02.003
LeDoux, J. E., & Pine, D. S. (2016). Using neuroscience to help understand fear and anxiety: A two-system framework. American Journal of Psychiatry, 173(11), 1083–1093. https://doi.org/10.1176/appi.ajp.2016.16030353
McEwen, B. S., Eiland, L., Hunter, R. G., & Miller, M. M. (2012). Stress and anxiety: Structural plasticity and epigenetic regulation as a consequence of stress.Neuropharmacology, 62(1), 3–12. https://doi.org/10.1016/j.neuropharm.2011.07.014
McEwen, B. S., Nasca, C., & Gray, J. D. (2016). Stress effects on neuronal structure: Hippocampus, amygdala, and prefrontal cortex. Neuropsychopharmacology, 41(1), 3–23. https://doi.org/10.1038/npp.2015.171
McEwen, B. S., & Wingfield, J. C. (2010). What is in a name? Integrating homeostasis, allostasis, and stress. Hormones and Behavior, 57(2), 105–111. https://doi.org/10.1016/j.yhbeh.2009.09.011
Nagasawa, M., Mitsui, S., En, S., Ohtani, N., Ohta, M., Sakuma, Y., … Kikusui, T. (2015). Oxytocin-gaze positive loop and the coevolution of human–dog bonds. Science, 348(6232), 333–336. https://doi.org/10.1126/science.1261022
Nesse, R. M., & Ellsworth, P. C. (2009). Evolution, emotions, and emotional disorders. American Psychologist, 64(2), 129–139. https://doi.org/10.1037/a0013503
Roozendaal, B., McEwen, B. S., & Chattarji, S. (2009). Stress, memory and the amygdala. Nature Reviews Neuroscience, 10(6), 423–433. https://doi.org/10.1038/nrn2651
Sundman, A.-S., Van Poucke, E., Svensson Holm, A.-C., Faresjö, Å., Theodorsson, E., Jensen, P., & Roth, L. S. V. (2020). Long-term stress levels are synchronized in dogs and their owners.Scientific Reports, 10(1), 17112. https://doi.org/10.1038/s41598-020-74204-8
Zovkic, I. B., Guzman-Karlsson, M. C., & Sweatt, J. D. (2013). Epigenetic regulation of memory formation and maintenance. Learning & Memory, 20(2), 61–74. https://doi.org/10.1101/lm.026575.112
Yawning is a simple, evolutionarily conserved behavior with physiological and social functions. In both humans and dogs, yawns can be contagious, reflecting motor and social resonance rather than necessarily empathic processes. Evidence indicates that dogs are more likely to yawn in response to familiar humans, particularly their owners, and that such yawning serves a pacifying and communicative function. Contagious yawning in dogs appears to facilitate social attunement, reduce tension, and signal peaceful intent. This paper reviews current research on canine yawning, its neural and behavioral underpinnings, and its role in interspecific communication, highlighting the ways dogs use yawning and related behaviors to maintain harmonious interactions with humans.
Your dog understands your yawn
A yawn is a simple behavior, a reflex with specific physiological functions. We are not the only ones yawning. Chimpanzees, bonobos, macaques, and dogs, among others, yawn (Guggisberg, Mathis, Schnider, & Hess, 2010; Joly-Mascheroni, Senju, & Shepherd, 2008; Ake & Kutsukake, 2023). Although a simple behavior, yawning also performs social functions. It is contagious not only within groups of individuals of the same species but also across species, including between humans and dogs (Joly-Mascheroni et al., 2008; Romero, Konno, & Hasegawa, 2013; Norscia & Palagi, 2011).
Because yawning is both widespread and multifunctional, several explanations have been proposed for its original biological role. One classic hypothesis suggests that yawning increases the influx of oxygen into the blood when carbon dioxide levels rise; however, this explanation is now widely considered unsupported (Guggisberg et al., 2010). Another hypothesis proposes that yawning stretches the muscles of the tongue and neck (Provine, 2012). A further interpretation emphasizes the need to maintain alertness, a crucial condition for predators (Provine, 2012). Given that social predators depend on one another, yawning may have evolved to be contagious through natural selection because of the cooperative advantages it confers. Additionally, yawning may help regulate brain temperature (Gallup & Gallup, 2007; Gallup & Eldakar, 2013; Gallup, 2022).
Pharmacological and neurochemical research shows that yawning is regulated by a network of neurotransmitters. Dopamine (via D₂/D₃ receptors) and serotonin both modulate yawning, and oxytocin may also play a role (Wani & Agarwal, 2025; Argiolas & Melis, 1998). These interactions suggest that yawning reflects changes in arousal, social state, and internal regulation—consistent with its role as a pacifying or self-soothing behaviour.
A widely proposed explanation for contagious yawning is that mirror-neuron systems in the frontal cortex of various vertebrates, including humans and dogs, activate corresponding motor representations in others. Neuroimaging studies in humans support this interpretation (Platek, Mohamed, & Gallup, 2005; Schürmann et al., 2005). Further neural evidence indicates that the ventromedial prefrontal cortex—a region associated with social processing—is also involved in contagious yawning, reinforcing the notion that the phenomenon is both motor-resonant and socially relevant (Nahab, Hattori, Saad, & Hallett, 2009).
Studies have found that dogs are more prone to yawn when their owners yawn than when strangers do (Romero et al., 2013; Silva, Bessa, & de Sousa, 2012). In the Tokyo study, researchers monitored the dogs’ heart rate and found no significant change across conditions, suggesting that the yawns were not merely a stress response (Romero et al., 2013). In one auditory-yawn study from Porto University, dogs yawned more to familiar than unfamiliar human yawns, and their stress-related behavior did not differ by condition—indicating that increased yawning was not simply a stress response (Silva et al., 2012). Meanwhile, an experiment at Birkbeck College (University of London) demonstrated that live human yawning triggers yawning in many dogs (Joly-Mascheroni et al., 2008).
Taken together, current behavioral evidence in dogs suggests that a form of interspecific resonance exists: dogs and humans can synchronize their actions during shared activities, and such coordination may emerge from motor-resonance mechanisms analogous to mirror-neuron systems (Lamontagne & Gaunet, 2024). Developmental evidence shows that contagious yawning in puppies emerges gradually, suggesting a maturational component to this resonance (Madsen & Persson, 2013). Mirror neurons may thus provide a neural basis not only for imitation but also for allelomimetic behavior.
However, whether contagious yawning reflects empathy remains debated. One recent Bayesian re-analysis of canine studies concluded that although contagious yawning is present in dogs, it does not display the familiarity, gender, or prosociality biases that an empathy model predicts (Neilands, Claessens, & Ren, 2020). Comparative research likewise cautions that contagious yawning cannot be taken as direct evidence of empathic capacity without more stringent criteria (Massen & Gallup, 2017).
Wolf yawning, a behavior shared by wolves and dogs and also common in other species (photo by Monty Sloan, Wolf Park, Indiana, USA).
The dog’s yawn is much like ours. It often precedes the same characteristic sound. While we commonly associate yawning with tiredness or boredom, it can also express embarrassment, insecurity, excitement, and relief. Some people even yawn when they’re in love—which, if misinterpreted, might be embarrassing.
Dogs may yawn when tired, but yawning usually serves a pacifying function, both for themselves and for others. As with many behaviors, what may have started as one function can evolve into others. Over time, yawning appears to have become a signal of peaceful intentions. For example, a male dog may yawn when a female snarls during courtship, signaling deference rather than aggression; or a confident dog may yawn at an insecure opponent to reassure it.
Dogs yawn at us with the same functions and results. They may also yawn as a displacement activity. An owner scolding his dog is a typical situation in which we see a dog yawn. In critical training cases prone to error, such as the so-called ‘stay,’ the owner’s behavior often causes the dog to feel insecure. A yawn is likely to follow, together with licking and muzzle-nudging. As soon as the owner changes behavior, say, by using a friendlier tone or more relaxed body posture, the dog ceases to display those pacifying behaviors.
Conclusion
Yawning is a ubiquitous behavior with ancient biological roots. While its original function remains debated, evidence supports multiple physiological and social roles—including thermoregulation, alertness maintenance, and behavioral synchronization. In dogs, as in humans, contagious yawning reflects a form of motor and social resonance, though not necessarily empathy in the strict scientific sense. Research consistently shows that dogs are more likely to yawn in response to familiar humans, particularly their owners, and such responses are not simply manifestations of stress. Rather, they appear to facilitate social attunement, reduce tension, and communicate peaceful intent.
Thus, when your dog yawns at you, it is unlikely to be random. It most likely expresses comfort and trust, and it invites the maintenance of social harmony. Your dog yawns at you to show it is friendly and peaceful—and you may safely yawn back, confirming the same. Yawning, along with champing (chomping), lip-licking, eye-squeezing, a pouty mouth, and the canine muzzle-grasp—all common elements of intraspecific canine social interaction—functions equally effectively in interspecific communication.
Featured Picture: Human and dog yawning (composition by Roger Abrantes).
References
Ake, K., & Kutsukake, N. (2023). Contagious yawning in African painted dogs (Lycaon pictus). Animal Cognition, 26(4), 1191–1198. https://doi.org/10.1007/s10071-023-01766-1
Gallup, A. C., & Eldakar, O. T. (2013). The thermoregulatory theory of yawning: What we know from over five years of research. Frontiers in Neuroscience, 6, Article 188. https://doi.org/10.3389/fnins.2012.00188
Gallup, G. G. Jr., & Gallup, A. C. (2007). Yawning as a brain-cooling mechanism: Nasal breathing and forehead cooling diminish the incidence of contagious yawning. Evolutionary Psychology, 5(1), 92–101. https://doi.org/10.1177/147470490700500109
Joly-Mascheroni, R. M., Senju, A., & Shepherd, A. J. (2008). Dogs catch human yawns. Biology Letters, 4(5), 446–448. https://doi.org/10.1098/rsbl.2008.0333
Lamontagne, A., & Gaunet, F. (2024). Behavioural synchronisation between dogs and humans: Unveiling interspecific motor resonance? Animals, 14(4), 548. https://doi.org/10.3390/ani14040548
Madsen, E. A., & Persson, T. (2013). Contagious yawning in domestic dog puppies (Canis lupus familiaris): The effect of ontogeny and emotional closeness on low-level imitation. Animal Cognition, 16(2), 233–240. https://doi.org/10.1007/s10071-012-0568-9
Massen, J. J. M., & Gallup, A. C. (2017). Why contagious yawning does not (yet) equate to empathy. Neuroscience & Biobehavioral Reviews, 80, 573–585. https://doi.org/10.1016/j.neubiorev.2017.07.006
Nahab, F. B., Hattori, N., Saad, Z. S., & Hallett, M. (2009). Contagious yawning and the frontal lobe: An fMRI study. Human Brain Mapping, 30(5), 1744–1751. https://doi.org/10.1002/hbm.20638
Neilands, P., Claessens, S., & Ren, I. (2020). Contagious yawning is not a signal of empathy: No evidence of familiarity, gender or prosociality biases in dogs. Proceedings of the Royal Society B: Biological Sciences, 287(1920), 20192236. https://doi.org/10.1098/rspb.2019.2236
Platek, S. M., Mohamed, F. B., & Gallup, G. G. Jr. (2005). Contagious yawning and the brain. Brain Research: Cognitive Brain Research, 23(2–3), 448–452. https://doi.org/10.1016/j.cogbrainres.2004.11.011
Provine, R. R. (2012). Curious behavior: Yawning, laughing, hiccupping, and beyond. Harvard University Press.
Romero, T., Konno, A., & Hasegawa, T. (2013). Familiarity bias and physiological responses in contagious yawning by dogs support link to empathy. PLOS ONE, 8(8), e71365. https://doi.org/10.1371/journal.pone.0071365
Schürmann, M., Hesse, M. D., Stephan, K. E., Saarela, M., Zilles, K., Hari, R., & Fink, G. R. (2005). Yearning to yawn: The neural basis of contagious yawning. NeuroImage, 24(4), 1260–1264. https://doi.org/10.1016/j.neuroimage.2004.10.022
Silva, K., Bessa, J., & de Sousa, L. (2012). Auditory contagious yawning in domestic dogs (Canis lupus familiaris): First evidence for social modulation. Animal Cognition, 15(4), 721–724. https://doi.org/10.1007/s10071-012-0473-2
Wani, P. D., & Agarwal, M. (2025). The science of yawning: Exploring its physiology, evolutionary role, and behavioral impact. Journal of Family Medicine and Primary Care, 14(8), 3115–3120. https://doi.org/10.4103/jfmpc.jfmpc_1677_24
Dogs also exhibit the muzzle grasp behavior (photo by Marco de Kloet).
A “Muzzle grasp” (or muzzle grab) is a common behavior shown by social canines, e.g., wolves (Canis lupus lupus), dingoes (Canis lupus dingo), and dogs (Canis lupus familiaris). The primary function of this behavior is to confirm a relationship rather than to settle a dispute. The more self-confident or higher-ranking individual will muzzle-grasp a more insecure or lower-ranking partner to assert its social position. The more insecure individual does not resist the grasp; on the contrary, it often displays submissive behavior, literally inviting its partner to muzzle-grasp it. Even though we sometimes see this behavior at the end of a dispute, wolves and dogs only use it toward individuals they know well—pack members—as a kind of saying, “You’re still a cub (pup).” The dispute itself tends not to be serious, merely a low-key challenge, often over access to a resource. Youngsters, cubs, and pups sometimes solicit adults to muzzle-grasp them. This behavior appears reassuring to them.
The muzzle-grasp behavior emerges early in development. Canine mothers muzzle-grasp their puppies (sometimes accompanied by a growl) to deter them from suckling during weaning. Field observations confirm this mechanism. As Packard, Mech, and Ream (1992, p. 1274) report, “In the context of playing, begging, and sharing, pups did not leave when another wolf muzzled, snapped, or lunged. In contrast, the muzzling by the nurser in the context of suckling terminated the pups’ attempts to gain access to nipples.” This observation illustrates the early communicative value of the muzzle contact as both a mild inhibitory and relational signal. Cubs and pups also muzzle-grasp one another during play, typically between six and nine weeks of age. They probably learn through play that the muzzle-grasp is an effective way of stopping an opponent from doing something, while also learning bite inhibition. If they bite too hard, they elicit a fight and risk injury. A muzzle-grasp, therefore, does not involve biting, only grasping. This behavior helps develop a relationship of trust between both parties—“we don’t hurt one another.”
Similar tactile interactions, including muzzle-to-muzzle contact, also occur in post-conflict and affiliative contexts among wolves. Cordoni and Palagi (2019) describe reciprocal muzzle-licking between adults and immature pack members following mild conflicts—acts that function as “consolation” and reinforce social bonds. Although a muzzle-grasp differs mechanically from muzzle-licking, both share an underlying functional value: the restoration or affirmation of trust within a dyad. These tactile gestures exemplify the nuanced physical vocabulary through which canids maintain cohesion and mitigate tension within the pack.
Classic naturalist observations (Zimen, 1981) describe frequent muzzle-to-muzzle contacts and note adults seizing pups’ muzzles during play and weaning; together with quantitative field data (Packard, Mech, & Ream, 1992), this supports the view that muzzle contact is an early-emerging, ritualised tactile signal rather than an aggressive act.
When used to settle a dispute, a muzzle-grasp may appear more violent and usually ends with the individual being muzzle-grasped exhibiting passive, submissive behavior. Yet participants very seldom, if ever, get hurt, an occurrence that would undermine the behavior’s function.
Left: Cubs and pups muzzle grasp one another during play. Right: Muzzle grasp in adult wolves (photos by Monty Sloan).
A muzzle-grasp requires self-control. Higher-ranking wolves and dogs muzzle-grasp their pack members (teammates) and, by doing so, confirm their rank while displaying restraint. Lower-ranking wolves and dogs often engage in muzzle-grasping behavior to affirm their social position and reassure themselves that they remain included in the group.
The muzzle-grasp behavior probably originated as both a form of maternal (and later paternal) control and as a play behavior among cubs. As it appears to have been beneficial to all parties involved, it may have become a factor favored by natural selection, spreading from generation to generation and evolving as any other trait that enhances the fitness of individuals within cohesive social groups.
In domestic dogs, when puppies are about five to seven weeks old, their mother regularly muzzle-grasps them to deter suckling. At first, her behavior frightens them, and they may whimper excessively, even though she does not harm them. Later, when grasped by the muzzle, the puppy immediately shows passive submissive behavior—lying on its back and exposing its ventral side. Previously, it was assumed that the mother needed to pin the puppy to the ground; however, Packard et al. (1992) observed that, in wolves, in practice, “[…] on the occasions when the nurser winced or muzzled the pups, the pups did not persist” and that “[…] counter-tactics for overcoming nurser rejection did not occur (pp. 1271–1272).” Most puppies submit voluntarily. Over time, this behavior pattern assumes variations. Wolf cubs and puppies often invite the alpha male (the leader of the pack and, in wolves, usually their father) as well as other adults to grasp them by the muzzle, thereby soliciting a demonstration of their elders’ superiority and self-control while simultaneously showing their own acceptance and submissiveness. This is among the most reassuring behaviors an adult can show a youngster.
Domestic dogs sometimes approach their owners puffing gently with their noses. By gently placing a hand around their muzzle, we may reassure them of acceptance, demonstrate self-control, and convey that they can trust us. That is speaking dog-language to the best of our abilities. After being muzzle-grasped for a while, the dog will usually show a nose-lick, perhaps yawn, and then walk calmly away. It is as if the dog were saying, “I’m still your puppy,” and the owner replied, “I know—and I’ll take good care of you.”
The muzzle-grasp behavior can be challenging to classify. Some researchers see it as social or affiliative, others as agonistic, and still others as pacifying. Because its primary function is to confirm and maintain relationships, it may best be considered a social behavior—a ritualized, low-intensity interaction that reinforces trust and cohesion within the group.
Next time your dog gently nudges or invites a muzzle‑grasp, pause for a moment—what you see as a simple dog behavior is, in canine language, a subtle conversation of trust and understanding.
References
Abrantes, R. (1987). Hundesprog. Borgen Forlag, Copenhagen.
Abrantes, R. (1997). The Evolution of Canine Social Behavior. Naperville, IL: Wakan Tanka Publishers.
Cordoni, G., & Palagi, E. (2019). Back to the future: A glance over wolf social behavior to understand dog–human relationship. Animals, 9(11), 991. https://doi.org/10.3390/ani9110991
Zimen, E. (1981). The wolf: His place in the natural world. Souvenir Press Ltd. ISBN 9780285624115
Note: I first wrote about the muzzle grasp behavior in canids in my Danish book Hundesprog (1987), where I called it “mund om snuden,” which translates directly as “mouth around the snout.” This term became “muzzle grasp” in the first English edition of the book, titled Dog Language. I later wrote Muzzle Grab Behavior in Canids on April 25, 2012. Two years afterward, on March 13, 2014, I revised it as Canine Muzzle Grasp Behavior—Advanced Dog Language. True to my philosophy of updating articles and papers as new evidence emerges, I have once again revised this work. The latest version, published in November 2025, appears here under the title Muzzle Grasp Behavior in Canids.
Scent detection has fascinated me since my early days as a student of biology, and I was already training detection animals at the beginning of the 1980s. Over the years, I have trained dogs, rats, and guinea pigs to detect narcotics, explosives, blood, vinyl, fungus, landmines, tuberculosis, and tobacco—and they excelled in all these tasks.
What has always intrigued me most is how deeply scent detection seems to be woven into their very being, regardless of species. Indeed, much before dogs became our partners in scent detection, olfaction had already shaped the mammalian brain—including ours. Although humans are often described as “microsmatic,” this view stems mainly from a 19th-century anthropocentric bias. In fact, human olfactory performance—when properly measured—can rival that of many other mammals (McGann, 2017). Fossil endocasts reveal that early mammalia forms possessed disproportionately large olfactory bulbs, suggesting that life for our distant ancestors was guided above all by smell (Rowe, Macrini, & Luo, 2011). The olfactory pathways remain among the most conserved in the mammalian nervous system, closely intertwined with limbic and reproductive circuits (Shipley & Ennis, 1996; Boehm, Zou, & Buck, 2005). As Lledo, Gheusi, and Vincent (2005) observed, “It is clear today that olfaction is a synthetic sense par excellence. It enables pattern learning, storage, recognition, tracking, or localization and attaches emotional and hedonic valence to these patterns” (p. 309). To smell, then, is not merely to detect—it is to think, feel, and remember.
Most of my detection work was carried out for the police, armed forces, SAR teams, or other professional agencies. Yet, I had written about scent detection already in the early 1980s, in my first book, Psychology rather than Force, published in Danish. Back in 1984, I called it “nose work” (a direct translation from the Danish næsearbejde). I recommended that all dog owners stimulate their dogs by giving them detection tasks, beginning with their daily rations. We even conducted some research on this, and the results were highly positive: dogs trained in detection work improved in many aspects of their otherwise problematic behavior. My recommendation remains the same today. Physical exercise is, of course, essential—but do not forget to stimulate your dog’s nose as well, perhaps its primary channel of information about the world.
Above: In “Hundesprog” (Dog Language) from 1987, I mention “nose work” with an illustration from Alce Rasmussen. To the right: Yours truly in 1984 with a Siberian Husky, an “untrainable” dog, as everybody used to say. This was when my book “Psychology rather than Force” created a stir. We were then right at the beginning of the animal training revolution. In that book, I mention “nose work” (a direct translation from the Danish “næsearbejde”) and recommend it as an excellent way to stimulate our dogs.
Recent field data illustrate how central olfaction is to the daily lives of canids. Wolves in the Białowieża Forest, for instance, were active on average 45.2 % of every 24 hours—about 10.8 h per day—primarily in movement, travelling, and search behaviours (Theuerkauf et al., 2003, Table 1, p. 247). Monthly patterns (Figure 6, p. 249) suggest that activity levels vary with season, although exact numerical ranges are not provided in the text. Comparable patterns appear in other canids: red foxes spend about 43 % of their observable foraging time sniffing the ground (Wooster et al., 2019), and free-ranging domestic dogs devote substantial portions of their active time to exploratory and searching behaviours—activities guided predominantly by olfaction (Banerjee & Bhadra, 2022). These figures reveal that for a wolf or fox, using the nose is not an occasional act but a continuous occupation, consuming many hours each day.
Measurement
%
Hours (h)
Time active
45.2 %
10.8
Time moving
35.9 %
8.6
Table 1. Average daily activity of wolves in the Białowieża Forest, Poland (1994–1999), showing the proportion of time spent active and moving, both as a percentage of the 24-hour day and in hours. Data from Theuerkauf et al. (2003, Table 1, p. 247).
Note. “Time active” includes periods when wolves were travelling, hunting, or otherwise moving. Observations indicate that these behaviours are predominantly guided by olfaction. Activity was generally higher at night, and seasonal variation appears linked to day length and prey availability. On average, wolves were active roughly half the day (~10.8 h), highlighting that extensive daily searching and tracking is a defining feature of their ecology (Theuerkauf et al., 2003, Table 1, p. 247).
When I began promoting “nose work” in the early 1980s, I did so from personal experience rather than data. I spent many hours on scent detection with my English Cocker Spaniels. They loved it and were calmer, more focused, and more fulfilled than their peers who were not as nose-stimulated. I quickly discovered that scent detection was so self-reinforcing—in behaviorist terms—that no other reinforcers were needed beyond my approval, which they actively sought. In those moments, I realised that to be a dog is to be a cooperative nose-worker.
Science has since validated that intuition.Scent work is not a modern invention—it is a structured expression of what canids have done for thousands of years: exploring their world through odor cues. When we engage a dog’s nose, we are not merely training a skill; we are restoring a function at the very core of its evolution. Understanding that is perhaps the greatest lesson of scent detection: to educate and enrich a dog’s life, we must first respect the sensory world in which it truly lives.
Boehm, U., Zou, Z., & Buck, L. B. (2005). Feedback loops link odor and pheromone signaling with reproduction. Cell, 123(4), 683–695. https://doi.org/10.1016/j.cell.2005.09.027
Lledo, P.-M., Gheusi, G., & Vincent, J.-D. (2005). Information processing in the mammalian olfactory system. Physiological Reviews, 85(1), 281–317. https://doi.org/10.1152/physrev.00008.2004
Rowe, T. B., Macrini, T. E., & Luo, Z.-X. (2011). Fossil evidence on origin of the mammalian brain. Science, 332(6032), 955–957. https://doi.org/10.1126/science.1203117
Theuerkauf, J., Kamler, J. F., & Jedrzejewski, W. (2003). Daily patterns and duration of wolf activity in the Białowieża Forest, Poland. Journal of Mammalogy, 84(1), 243–253. https://ibs.bialowieza.pl/publications/1396.pdf
Wooster, E., Wallach, A. D., & Ramp, D. (2019). The Wily and Courageous Red Fox: Behavioural analysis of a mesopredator at resource points shared by an apex predator.Animals, 9(11), 907. https://doi.org/10.3390/ani9110907
Featured image: Springer Spaniel, nose down, focused on a search.
Note: This article is a substantially revised and edited version of an earlier article from May 6, 2014, entitled Do You Like Canine Scent Detection? The revisions are extensive enough that the article deserves a new title and is therefore republished as new.
It dawned on me the other day, while at sea, one of those days with scattered clouds on the horizon and a fair wind barely sufficient to keep the boat sailing. Simplicity, that’s what makes it so soothing and scaringly beautiful. The sea invites you to dream, but make no promises; it is what it is, neither more or less. Be wise, and it will reward you; be foolish, and it will punish you.
You can’t hide at sea; you’ll encounter yourself whether you want it or not. The only viable strategy is honesty and integrity. It’s all so simple. The sea possesses this power, I discovered—the pertinent appears suddenly as frivolous, and the complex reveals itself in all its simple parts.
I felt absolutely ecstatic, like something major was happening, yet nothing particular stood out. As far as the eye could see, the world was an endless expense of blue, only slightly interrupted by a thin line, far, far away. Sea and sky, a few clouds on the horizon, the sun to the west, no birds, no fish, no sounds but the slight, rhythmic splashes of the boat gracefully cutting through the water, almost as silently as the flight of the owl.
Simplicity—I suppose, is what fascinates me most about Darwin’s brilliant concept of evolution by means of natural selection. The algorithm the survival of the fittest is the simplest idea one can conceive, and yet so powerful that it cuts through everything our understanding touches.
I have come to view the principle of simplicity as an old friend, always by my side as long as I can remember. From my young student days to the times of book writing or when on practical commissions, my friend Simplicity has been there, unobtrusively muttering, “Seek the simple…”
Theprinciple of simplicity, as such, was first propounded by the English philosopher, William of Occam (1300-1349). We also know it as Occam’s Razor: “Entia non sunt multiplicanda praeter necessitatem,” which is Latin for “Entities should not be multiplied more than necessary,” or “If two assumptions seem to be equally valid, the simpler one should be preferred.”
Simple is beautiful and simpler is beautifuller—and the sea has this influence on you. Thus, I took the liberty to apply the principle of simplicity to its own definition, and three corollaries emerged.
“If you have more than one option, choose the simplest.”
First corollary: “If you have only one option, you don’t have a problem; don’t waste your time complaining, just accept it and keep smiling!”
Second corollary: “If you don’t like having only one option, work to create more; then you’ll have the problem of choosing one.”
Third corollary: “If you don’t like having problems, don’t create options.” Return, then, to the first corollary, don’t complain, and keep smiling!
And so it is that I continue sailing across this vast sea of blue, feeling my heart beating for every, ever-so-gentle splash of the hull against the water. It all seems so simple: I am just a little ripple in the immense ocean, yet I am alive, and hence I must embrace life fully for as long as I can.
________________
Featured image: A few clouds on the horizon and a fair wind, barely enough to keep the boat sailing.
Note: This blog is an updated version of the original post “I’m Alive and I Have Only One Option” published on April 21, 2014. I made minor adjustments to the language to better convey my thoughts, for I had struggled with a few sentences in the original. The text is now more to my liking; however, upon re-reading it, I noticed a slight but significant change in its undertone, which is why I felt it deserved a new title. And yes, I kept the term “beautifuller.” It’s archaic from the 1800s, I know, but since I’m getting pretty archaic myself, I feel it’s fitting.
Walsh, D. (1979). Occam’s razor: A principle of intellectual elegance. American Philosophical Quarterly, 16(3), 241–244. https://www.jstor.org/stable/20009764.
—Do We Think Before We Act, or Do We Act Before We Think?
“Freedom is the individual’s capacity to know that he is the determined one, to pause between stimulus and response and thus to throw his weight, however slight it may be, on the side of one particular response among several possible ones.” This quote is from page 103 (of which I give you a photo below) of Lloyd-Jones, E., Westervelt, E. M. (1963) Behavioral science and guidance: proposals and perspectives. New York: Bureau of Publications, Teachers College, Columbia University.
We find this text frequently misattributed, misquoted, modified, and adulterated to suit various purposes and agendas on today’s overwhelming social media, where everybody knows everything—and nobody knows nothing (as they say).
Let’s take a moment to ponder and analyze what’s behind the (original) quote, without the social media razzle-dazzle.
There is a space—a tangible time gap—between the moment of an event and our reaction that we might use productively if we intentionally allow it to be there and engage it critically. That is, in other words, what the author means. It is an enticing proposition, grounded in solid research from previous decades.
The question of free will is a millennia-long one, in which neurophysiology only recently entered the fray. Theories of free will focus on two fundamental questions: its possibility and nature. Some define free will as the capacity to make choices undetermined by past events. Determinism, on the other hand, sustains that only one sequence of events is possible and is inconsistent with free will. In contrast, compatibilism maintains that free will is consistent with determinism.
For over twenty years, experiments have suggested that, unbeknownst to us, a substantial part of mental processing occurs unconsciously, i.e., even before we know we plan to act. When we become aware of the brain’s actions, we ponder and mistakenly believe our intentions have caused them. “Your decisions are strongly prepared by brain activity. By the time consciousness kicks in, most of the work has already been done,” according to Haynes (2013).
Many brain processes occur automatically and without involving our consciousness. That is a defensive mechanism preventing our minds from being overloaded by basic routine chores.
We assume our conscious mind makes decisions. Our current findings question this. Researchers—using fMRI brain scans—could predict participants’ decisions up to seven seconds before the subjects had consciously made them.
If we decide before we are even aware of it, then the question is what mechanism decides for us. The prevailing view in neuroscience is that consciousness is an emergent neural phenomenon. The firing of the brain’s neurons gives rise to consciousness and the sensation of free will or intentional action. These findings (Libet 1985) may not surprise neuroscientists who believe consciousness arises from brain activity (rather than brain activity originating from consciousness) since they view the conscious experience of free will as an emergent phenomenon of brain activity.
Researchers conclude that cerebral initiation of a spontaneous voluntary act begins unconsciously. However, within about 150 ms after the precise, conscious purpose emerges, we can still deliberately control the ultimate decision to act. Subjects can “veto” motor function for roughly 100–200 ms before a set time to act. That is the gap, the pause, so quoted and misquoted.
Whether that gap suffices to overcome the centuries-old free will quandary is highly arguable.
________________
References
Dennett, DC (2003). Freedom Evolves. Viking Press.
Fischer, JM & Ravizza, M (1998). Responsibility and Control: A Theory of Moral Responsibility. Cambridge University Press.
Haynes, J-D (2013). World.Minds: Do We Have Free Will? (Charité Berlin).
Libet, B (1985). Unconscious cerebral initiative and the role of conscious will involuntary action. Behavioral and Brain Sciences,8(4), 529-539. doi:10.1017/S0140525X00044903.
Park, H-D et al. (2020). Breathing is coupled with voluntary action and the cortical readiness potential. Nature Communications, 2020; 11 (1) DOI: 10.1038/s41467-019-13967-9.
Pereboom, D (2001). Living Without Free Will. Cambridge University Press.
Soon, C, Brass, M, Heinze, HJ, et al. (2008). Unconscious determinants of free decisions in the human brain. Nat Neurosci11, 543–545. https://doi.org/10.1038/nn.2112
Soon, CS, He, AX, Bode, S, Haynes, J-D (2013). Decoding abstract intentions, Proceedings of the National Academy of Sciences. March.
Strawson, GJ (1994). The impossibility of moral responsibility. Philosophical Studies 75 (1-2):5-24.
___________________
Photo: Painting “The Illusion of Free Will (2016) by Daniel Stroup.
Your comments are welcome. Please feel free to leave a reply. Cancel reply
The author, Roger Abrantes, and Petrine, English Cocker Spaniel, in 1984.
“A reinforcer is not a reward.”
Some things must be said again—and again.
For the umpteenth time, a reinforcer is not a reward. When I hear “Force-Free” trainers say, “dogs like to work to earn rewards,”1 I suspect and fear they miss by a mile and a half the essence and function of reinforcers in learning theory (and so also of inhibitors).
I’m not splitting hairs. There is a crucial difference between reinforcing a particular behavior and rewarding an individual. I suspect ignorance hereof is also the cause of the many incorrect statements on inhibitors2 from the “Radical Force-Free” camp (emphasis on radical).3
When terminology goes wrong
Look, I shouldn’t care less because reinforcers and the like are behaviorist things, and I’m an ethologist, not a behaviorist. But I do care, because my mind becomes strangely uneasy whenever I hear or see something fundamentally flawed and inconsistent.
If you claim the mantle of behaviorism, the least you can do is use its language correctly.
If you’re a behaviorist—and that’s what trainers in the “Radical Force-Free” camp are supposed to be—then at least be fair to the founding fathers of your learning philosophy and use the right terminology. All the rest seems to me an affront, disrespectful, and proof of unforgivable ignorance. Forgive the bluntness, but someone had to say it.
Back to the 1980s
“Dogs like to work to earn rewards” reminds me of the 1980s, when I opposed the old-school, military-style approach to dog training. The classes had all the charm of a drill parade: straight lines, sharp commands, leash jerks, and very little interest in what the dogs themselves thought of the matter—or whether they understood what we wanted of them. After ten minutes of marching back and forth, the instructor would say:
“And halt! Now, praise your dogs.”
Yes, I went to that kind of dog training with the first dog I had as an adult. That was the training we had back then.
I walked out in disgust with the sort of calm determination only youthful indignation can fuel. I decided there and then that Petrine, my dog, and I would train on our own and show them. We did.
Discovering training through ethology
I substituted praise with reinforcers—the real thing, not rewards—including my dygtig⁴ and a few treats given at strategic times and points. I stopped using a leash and started using a lead. Again, not splitting hairs—it makes a huge difference what you think of, and how you use, that piece of rope or leather that connects you to your dog.
A leash leashes; a lead leads. It’s as simple as that.
Leash jerks gave way to “No,” immediately followed by “dygtig,” when Petrine, not me, corrected the mistake. She seemed almost pleased to catch her own slips, as if this strange little team sport of ours finally made sense. She visibly enjoyed being my “teammate,” a role she took with disarming seriousness.
Learning from the giants
I was, then, a student of ethology, and I knew about social animals and social canines, including our domestic dog, and how contact, social acceptance, and feeling safe functioned as unconditioned, primary reinforcers (‘benefits of group living’ in ethology jargon). The social canines were among my favorites; I studied them diligently, inspired by my fellow senior students: Eberhard Trumler, Erich Klinghammer, Thomas Althaus, and Erik Zimen—alas, all gone now.
Old, venerable Professor Lorenz’s words rang in our ears by then (they still do):
“To understand an animal, first you have to become a partner.”
And so I did, applying to my training the best principles of ethology that I had learned from the great teachers—Lorenz, Tinbergen, von Frisch—and from my prominent elder colleagues. Later, I would even incorporate selected elements of behaviorism into what eventually became Animal Training My Way, but that is another story for another time.
Winning where no one expected us to
At the end of the term, I signed up for the final “obedience” competition at the club, a hunting dog club run by real green-clad hunters, and we won with max points. That a young long-haired fellow in faded Levi’s and clogs had won created some agitation—and to add insult to injury, my dog was a little, red, seven-month-old English Cocker Spaniel (a genuine one, not one of those oddballs we see in the US today), female on top of all.
Petrine was intelligent, beautiful, charming, a workaholic, and a sweetie-pie—though I suppose I was already helplessly devoted to her.
Our performance raised eyebrows and drew more humming than the establishment would have wished. At the prizes-and-punch social function, a few civilians asked me in a whisper whether I would help train their fidos (read: companion dogs).
And just like that, I was in deeper than I thought
The following Saturday, we were training on a grassy field across the road from where I lived, which is now the local firemen’s station. That was 1982, the summer before my son Daniel was born, and that’s how dog training came into my life. I never planned it.
Two years later, in 1984, I wrote my first book, Psychology Rather than Force, with far too little experience but loads of good ideas, including force-free, hands-free, reinforcement-based training—alas, all terms used as slogans these days—with as few inhibitors as possible; and it even included a whistle (the precursor of the clicker).
I was positive dog training would change. It did, and the rest is history.
And if this story has a punch line, it is this: reinforcers do the work—rewards are just the icing.
Notes
This is an actual quote from a document published online by a confessed “Force-Free” trainer. Note that Skinner writes about reinforcers and rewards, “The strengthening effect is missed, by the way, when reinforcers are called rewards. People are rewarded, but behavior is reinforced. If, as you walk along the street, you look down and find some money, and if money is reinforcing, you will tend to look down again for some time, but we should not say that you were rewarded for looking down. As the history of the word shows, reward implies compensation, something that offsets a sacrifice or loss, if only the expenditure of effort. We give heroes medals, students degrees, and famous people prizes, but those rewards are not directly contingent on what they have done, and it is generally felt that the rewards would not be deserved if they had been worked for” (Skinner, 1986, p. 569). ↩︎
In 2013, I suggested we change punisher and derivatives to inhibitor and derivatives to avoid the moral and religious connotations of the former, particularly in Latin languages, and to emphasize their function and use as a learning tool. ↩︎
“Radical Force-Free dog trainers” is my denomination for those trainers adhering to the positive or force-free movement, but having extreme views like claiming positive reinforcers are the only learning tool one needs, they never use aversive stimuli, one should never say “no,” everyone else but them is wrong, and other absurdities. Please do not confuse with the non-radical positive, force-free dog trainers who are equally force-free but sensible, open-minded, prudent in their claims, and polite and considerate to other thinkers. ↩︎
Reference List (First Editions in Original Language)
A tribute to my great teachers and senior fellow students:
Althaus, T. (1982). Verhaltensontogenese beim Siberian Husky [Dissertation, Universität Bern]. Institut für Zoologie.
Klinghammer, E. (Hrsg.). (1979). The behavior and ecology of wolves. Garland STPM Press. (Original language: English; this edited volume was first published in English.)
Lorenz, K. (1949). Er redete mit dem Vieh, den Vögeln und den Fischen. Borotha-Schoeler. (First German edition; this is the original form of what later became known in English as King Solomon’s Ring.)
Tinbergen, N. (1948). De natuur van het dier. Het Spectrum. (First Dutch edition; predates the 1951 English The Study of Instinct.)
Trumler, E. (1961). Der Hund. Georg Müller Verlag. (His earliest and most influential dog-ethology book; later expanded works followed.)
von Frisch, K. (1927). Aus dem Leben der Bienen. Springer. (First German edition; foundational to his later Tanzsprache works.)
Zimen, E. (1971). Wolfsfibel. Kosmos Gesellschaft der Naturfreunde. (Zimen’s first book-length publication in German; precedes later major works such as Der Wolf.)
References mentioned in the blog
Abrantes, R. (1984). Psykologi Fremfor Magt (Psychology Rather Than Force). Lupus Forlag.
Abrantes, R. (2015). Animal Training My Way. Wakan Tanka Publishers.
Abrantes. R. (2013). The 20 Principles All Animal Trainers Must Know. Wakan Tanka Publishers.
Featured photo: Roger Abrantes and Petrine in 1984 by Annemarie Abrantes.
📦 Glossary
Reinforcer
A consequence that increases the likelihood of a behavior being repeated. In learning theory, a reinforcer is defined functionally, not emotionally: it works because it strengthens behavior, not because it feels like a “reward.”
Reward
A colloquial, subjective term for something the giver believes is pleasant to the recipient. Unlike reinforcers, rewards do not have to change behavior—and often don’t. This is why “reward” ≠ “reinforcer.”
Inhibitor
A consequence that reduces the likelihood that a behavior will recur. The functional opposite of a reinforcer. Inhibitors are not “punishment” in the everyday sense—they can be as subtle as social disengagement or loss of access.
Dygtig(Danish)
Pronounced roughly “Dö-gtee.” Literally “skilled” or “good,” introduced as a conditioned (or semi-conditioned)reinforcer in training (Abrantes, 1984). The “dygtig,” delivered with timing and consistency, and a friendly facial expression/body language, signals to the dog: “That was correct—keep doing it.” It embodies applied ethology at its best.
Lead vs. Leash
Lead: A tool intended to guide the dog; used with communication in mind. Leash: A tool that often defaults to restraint. The distinction reflects mindset more than equipment—a leash leashes; a lead leads.
Force-Free (in practice)
A training approach aiming to avoid aversive physical force. Initially grounded in learning theory, but currently used in ways that often blur terminology and introduce inconsistencies between science and practice.
Primary Reinforcer
A stimulus that is naturally reinforcing without learning (conditioning)—for example, food, social contact, safety, and touch (in most circumstances).
Conditioned Reinforcer
A neutral stimulus (e.g., dygtig, a whistle, a click) that becomes reinforcing through association with a primary reinforcer.
“I didn’t fail. I discovered 34 ways that don’t work!” I told them. But allow me to start from the beginning.
Success and failure are not absolutes; they are states of mind—criteria we adopt, perspectives we choose. Success may boost confidence and smooth the path ahead. Failure, conversely, fortifies resolve and builds persistence. Moving from one triumph to the next feels comfortable—perhaps too comfortable. Moving from one setback to another is taxing. So which is better? Let me tell you a story.
Once—young, spirited, and incapable of declining a challenge—I accepted a job trailering a horse. At the time, horse trailering was the number-one complaint among horse owners, much as home-alone issues plague dog owners today, and inappropriate elimination troubles cat owners.
Failing to trailer a horse at home is inconvenient; failing when you are hundreds of miles away is a genuine predicament. These owners were 200 miles from home. Their mare had refused to load after an equestrian event. They tried everything they knew, and everything others had told them, and still failed. Exhausted, they left the horse in a stall and drove home—then called me, offering anything if I could bring the mare back.
In hindsight, after hearing how many seasoned horsemen had tried—and how they had tried—I probably should have declined. But youth thrives on challenges, and so I drove to meet the horse.
We released her into a medium-sized arena, backed the trailer in, and I sat on the fence observing. She was a beautiful four-year-old quarter horse mix—alert, sensitive, and expressive. The owners recounted her history: no issues except trailering. They succeeded perhaps one in twenty attempts, and only after considerable distress. It was getting worse.
I will spare you the long list of misguided attempts employed before my arrival—well-intentioned efforts born more of frustration and inherited habits than of horsemanship. Don’t get me wrong: neither the owners, who were friendly and educated, nor those who tried to help were ill-disposed; they were simply relying on long-standing traditions that were not always gentle and had rarely been questioned.
These days, dog people often spend considerable time passionately disagreeing over training details. I sometimes invite them to visit the horse world, where perspective comes swiftly. Faced with the challenges still common in horse training, many of these canine disputes appear trivial. With a few admirable exceptions—brave horsemen and women who work to demonstrate that there are other, equally (or more) effective ways of handling a horse than sheer force—the field of horse training has long struggled to move beyond methods that rely mainly on force.
I stepped into the arena bare-handed—not even a rope. I liked the mare immediately. As with people, some animals evoke instant affinity; others do not. She seemed comfortable with me, too, if not from the beginning, then soon after. We walked quietly around the arena, each minding our own business. The owners left to run errands, a relief to both of us, I suspect.
After some time, the mare approached. We paused, inhaling and exhaling deeply. Horses do this when meeting. When in Rome, do as the Romans do;1 with horses, I become as equine as I reasonably can. It may look peculiar to some, but it works, and it matters. The more I attune myself to the animal’s own signals, the more she can evaluate me on her terms rather than mine.
We spent about two hours simply walking together. At times, she followed me slowly, three to four meters behind. At other moments—particularly when we passed a section of the fence where buckets and gardening tools were stored outside—she hesitated. Each time, I waited and behaved as though nothing were amiss. After a while, she would approach, and after six or seven such encounters, she passed the “scary” spot without difficulty. The same pattern emerged near the trailer.
Occasionally, I stopped, leaned against the fence or the trailer, and looked at her. She looked back and then approached slowly until she stood about a meter in front of me, relaxing into that characteristic stance with one hind hoof resting. Gradually, she began to follow me without hesitation. She had discovered, on her own terms, that I posed no threat and was sufficiently trustworthy.
When the owners returned, they asked whether I had loaded her.
“No,” I said.
“Oh, we’re sorry you failed,” they replied, offering polite condolences.
“Oh no,” I said. “I didn’t fail. I discovered 34 ways that don’t work to load a horse.” Strictly speaking, that was not entirely true. I had made a few tentative attempts early on to see how she reacted to the trailer, but I had not tried to load her in earnest. Most of the time was spent observing—learning what unsettled her, what reassured her, and how she chose to follow when she felt safe. There were easily thirty-four or more such small trials, each one teaching me something about her thresholds and preferences. What looked like inaction was, in fact, the slow process of letting the horse teach me how to proceed.
A few hours later, both the mare and I were standing inside the trailer, eating carrots and breathing calmly together. I never used a rope. I never used the halter. I never touched her until we had already been inside the trailer for a while.
So what happened? In short, applied ethology happened. I relied on the horse’s natural need for companionship and safety, and I tried to provide precisely that. I let her approach in her own time, assess my intentions, and decide whether to come closer. In behavioristic terms, the comfort of social proximity reinforced her tendency to follow me. Falling behind or moving away were aversive experiences that briefly left her feeling exposed, prompting her to return. But the crucial point is that she controlled the reinforcers and inhibitors2herself, not I.
That insight has shaped all my work with animals, no matter the species, though I hesitate to call it “training,” for what I value is the interaction—the relationship, the shared routines that work for both human and animal. It is a givers/takers game. Both partners must contribute, both must give and take—give what you can, take what you need.
The magic ingredient, I discovered, is humility, though not submission; confidence, though not dominance; being present as a trustworthy guide who points out possibilities and offers choices, while remaining as unobtrusive as possible in the background. It is, in truth, a humbling experience to realize how much one can learn, and how much better one can become, simply by listening to a horse.
Once she, the mare, had reliably chosen my company, walking into the trailer was merely the next natural step. After passing the trailer several times together, I entered it casually—no theatrics, no pressure, no fireworks either, once done. She followed without hesitation.
That day, I learned how to trailer a horse—not because I succeeded after four hours, but because I had uncovered 34 or so approaches that did not work. I thought I knew the procedure already; I was mistaken. Early success had lulled me into complacency.
Success and failure are not properties of events but of perspective. In the end, success came not from doing more, but from doing less—listening more carefully, and allowing the horse to teach me what I needed to know.
Notes
The expression “When in Rome, do as the Romans do” (Latin: Si fueris Romae, ieiunato sicut Romanum est) is attributed to Saint Ambrose (4th century). When Augustine asked why the Milanese practice differed from that of Rome, Ambrose replied, “When I am in Milan, I do not fast on Saturday; when I am in Rome, I fast on Saturday.” Hence the proverb, meaning that one should adapt to local customs. The practice of adapting and imitating also serves us well when studying and interacting with other species, as Konrad Lorenz and Jane Goodall, among others, have demonstrated. ↩︎
A reinforcerreinforces—meaning it strengthens behavior (it increases; one gets more). This is intuitive and consistent with standard usage. Likewise, an inhibitorinhibits—meaning it weakens behavior (it decreases; one gets less)—and the term explicitly states this functional effect. By contrast, the technical and classic term punisher refers to a stimulus that also weakens behavior, yet the word itself does not convey that effect. One may “punish” in the everyday sense without actually decreasing the behavior if the intensity or timing is inappropriate. The term inhibitor avoids this ambiguity: by name and by definition, it inhibits, that is, it weakens the behavior—without implying any difference in the underlying behavioral process. It also focuses purely on the functional outcome—a decrease in behavior rate—stripping away the ethical and emotional connotations associated with punishment. I am merely clarifying a conceptual point, not expecting that the established terminology will change. Ultimately, the choice of term is a matter of preference, and whichever term you use will require a clear definition for your readers and for anyone with whom you discuss these concepts. ↩︎
References
Budiansky, S. (1997). The nature of horses: Exploring equine evolution, intelligence and behavior. Free Press. ISBN 9780684827681.
Hempfling, K. F. (1993). Dancing with horses: The art of body language. Trafalgar Square Publishing. ISBN 9781570761517.
Parelli, P., & Parelli, K. (1993). Horsemanship: Theory & horse behavior. Parelli Natural Horsemanship. ISBN 9780965853300.
Roberts, M. (1997). The man who listens to horses: The story of a real-life horse whisperer who revolutionized the way we communicate with horses. Random House. ISBN 9780679456582.
Sinclair, E. (Director). (2016). Taming wild: A girl and a mustang [Film]. Taming Wild Productions.
Featured image: After some time, the mare approached. We paused, inhaling and exhaling deeply. Horses do this when meeting (photo from the EI files).
This article is an expanded and substantially updated version of the original “Animal Training—I Didn’t Fail, I Discovered 34 Ways That Don’t Work!” from June 4, 2014, which is why it is now published under a new title.
Canine maternal behavior is more than just feeding the pups. It is also to teach them dog language (Illustration by Alice Rasmussen from “Dog Language” by Roger Abrantes).
Watching dog mothers take care of their pups continues to fascinate me, and the large populations of village dogs in Africa and Thailand, where I spent and spend a great deal of my time, provides me with plenty of opportunities to do it. Village dogs are domestic dogs, not wild dogs. Often classified as stray dogs by the inept, ignorant eye of the western tourist, these dogs perform an important task in their communities of humans and their domestic animals.
Maternal behavior is behavior shown by a mother toward her offspring. In most species, it is the mother that primarily takes care of the youngsters, and the dog is no exception. Natural selection has favored the evolution of this particular behavior of the females.
In wild canids, although it is mostly the female that takes care of the puppies, the father (also called the alpha male) and other adults do become interested in the feeding and raising of the puppies when they begin emerging from the den. In the studies my team did in the 80s, our dogs showed the same pattern in a domestic set-up.
Maternal behavior is, thus, almost identical in wild a canids and domestic dogs. Immediately after birth, the mother dries the puppies, keeps them warm, feeds them and licks them clean. The maternal behavior right after birth is controlled by hormonal processes and problems may occur if the female gives birth too early. On the other hand, pseudo-pregnancy causes females to undergo hormonal changes which may elicit maternal behavior in various degrees. Maternal behavior seems to be self-reinforcing. Studies show that the levels of dopamine increase in the nucleus accumbens (a region of the brain) when a female displays maternal behavior.
When the puppies become older, the mother begins to educate them. She gives them the first lessons in dog language about the time weaning begins. Growling, snarling and the various pacifying behaviors are inborn, but the puppies need to learn their function.
The canine mother has three main tasks: (1) to feed the puppies, first with her own milk, then by regurgitation, (2) to keep them clean and warm, especially when they are very young, and (3) to educate the puppies.
A good canine mother is patient and diligent. When the puppies grow, dog owners often misunderstand the mother’s apparently more violent educational methods. She may growl at them and even attack them, but she never harms them. Muzzle grabbing (see illustrations) is fairly common. Without the mother’s intervention, the puppies would never become capable social animals and would not be able to function properly in a pack (a group of wild dogs living together is in English called a pack). When the puppies are about 8-10 weeks old, the mother seems to lose some of her earlier interest in them. In normal circumstances, the rest of the pack, then, takes over the continuing education of the puppies, their social integration in the group (which probably mostly consists of relatives) and their protection.
Dog owners sometimes report problems, e.g. that the mother has no interest in her puppies, or is too violent towards them. These problems are mainly due to our selective breeding (we select for beauty and utility while nature selects for overall fitness, hereby included adequate maternal behavior) and to our lack of understanding of the mother’s needs during and after birth, which often result in the female showing stress, insecurity or aggressive behavior.
Maternal effect is the mother’s influence on her puppies. It can have such an impact on certain behavior patterns that it can be difficult to distinguish between maternal effect and the effect of genetics. For example, observations have shown that a female reacting too nervously or fearfully toward certain sounds may affect her puppies into developing sound phobias beyond what we would expect given the puppies’ specific genotype. The strong influence of the maternal effect on the behavior of her puppies is the main reason why it is extremely difficult, if not impossible, to assess the hereditary coefficient for particular traits.
Bottom-line: Do not breed females that you suspect will not show reliable maternal behavior. Do not disturb a female with her pups more than absolutely necessary. A good canine mother knows better than you what’s best for her pups.
Fox, M. 1972. Behaviour of Wolves, Dogs, and Related Canids. Harper and Row.
Lopez, Barry H. (1978). Of Wolves and Men. J. M. Dent and Sons Limited.
Mech, L. D. 1970. The wolf: the ecology and behavior of an endangered species. Doubleday Publishing Co., New York.
Mech, L. David (1981). The Wolf: The Ecology and Behaviour of an Endangered Species. University of Minnesota Press.
Mech, L. D. 1988. The arctic wolf: living with the pack. Voyageur Press, Stillwater, Minn.
Mech. L. D. and Boitani, L. 2003. Wolves: Behavior, Ecology, and Conservation. University of Chicago Press.
Scott, J. P. and Fuller, J. L. 1998. Genetics and the Social Behavior of the Dog. University of Chicago Press.
Zimen, E. 1975. Social dynamics of the wolf pack. In The wild canids: their systematics, behavioral ecology and evolution. Edited by M. W. Fox. Van Nostrand Reinhold Co., New York. pp. 336-368.
Zimen, E. 1982. A wolf pack sociogram. In Wolves of the world. Edited by F. H. Harrington, and P. C. Paquet. Noyes Publishers, Park Ridge, NJ.
Do dogs see colors? Does that affect our dog training in any way?
In the early 1980s, we conducted tests at the Ethology Institute to determine whether dogs were colorblind, as popular opinion held. The conclusion of our experiments was that they could distinguish some colors but could not discriminate others. They were not completely color blind (seeing only shades of gray). They were more like some people who see colors, though not the whole spectrum. However, at the time, we could not determine whether the dogs’ color discrimination was due to distinguishing between real colors or to distinguishing between various shades of gray. Meanwhile, more modern research has cast some light on these questions.
Eyes contain light-catching cells (cones) that respond to color. Canines have fewer cones than humans, which implies that, in principle, their color vision cannot be as good as ours. To see colors, we need different types of cones that detect different wavelengths of light. We have three types of cones, which allow us to register the full range of color vision. Dogs, in contrast, have only two types of cones.
Researchers at the University of California, Santa Barbara, tested dogs’ color vision in the late 1980s. Their studies confirm that dogs see color, though not as well differentiated as humans do. For us, the rainbow looks violet, blue, blue-green, green, yellow, orange, and red. For a dog, we presume it looks dark blue, light blue, gray, light yellow, darker yellow, and very dark gray. Violet and blue are both perceived largely within the blue range (Neitz, Geist, & Jacobs, 1989).
Studies performed by Russian scientists demonstrated that dogs tend to discriminate real color rather than brightness cues (Kasparson, Badridze & Maximov, 2013). They match any color they register with no more than two pure spectral lights.
Dogs are dichromatic, as are most placental mammals. The ability to see long wavelengths necessary to distinguish red from green seems to have disappeared during evolution, probably during early mammalian evolution in the Mesozoic, likely under strong nocturnal selection pressures (Jacobs, 2009). Dichromatic vision, though, is good at distinguishing colors in dim light, favoring the most nocturnal animals.
Trichromats, like most humans, have three color-detecting cones (blue, green, and red) and can distinguish a vastly expanded range of color discriminations compared with dichromats. Trichromacy in humans is not ancestral to mammals. It is a derived re-evolution in Old World primates, an adaptation to a foraging ecology in which red–green discrimination provides a direct foraging advantage (Dulai et al., 1999). So human trichromacy is an exception produced by a very specific ecological niche, not the mammalian norm. Humans are visual outliers among mammals. When we treat human color perception as ‘normal,’ we are committing a deep phylogenetic bias.
Similar ecological pressures have shaped trichromatic vision independently in other, very distant lineages. The honeybee, Apis mellifera, is also trichromatic, seeing ultraviolet, blue, and green instead of blue, green, and red. Honeybee trichromacy is an adaptation shaped by the coevolution of pollinators and flowering plants, as many floral signals are specifically tuned to ultraviolet patterns invisible to human vision (Peitsch et al., 1992).
Human = A and C. Dog = B and D. It is difficult for the dog to discriminate between red and green.
The term “colorblind” is therefore misleading. Dogs are not deficient humans—they retain the ancestral mammalian visual condition. Some animals developed the ability to see certain colors, and others to see others, all depending on ecological pressures, mutations, and the subsequent costs and benefits each strategy implied for their struggle for survival (Jacobs, 1993).
What does this mean for our communication and training of our dogs? Since dogs find it difficult to distinguish between certain reds and greens (like some humans do), we should choose toys and training aids in other colors. For example, light blue or yellow are much easier colors for a dog to detect (Neitz, Geist, & Jacobs, 1989). On the other hand, when training them in any scent-detection discipline (Gazit & Terkel, 2003; Horowitz, 2009), we should use colors for targets that are difficult for them to see to compel them to use their noses rather than their eyes.
Training an animal entails altering its behavior, a process that can profoundly affect the individual and, if misapplied, lead to unintended and undesirable side effects. It is therefore sound practice to examine how the animal perceives and interacts with its world, as well as the evolutionary mechanisms that have shaped both the species and the individual under consideration. In other words, we should strive to see the world through the animal’s eyes—however imperfect that vision of ours may be. Only then can training be both effective and respectful for the animal and for us.
References
Dulai, K. S., von Dornum, M., Mollon, J. D., & Hunt, D. M. (1999). The evolution of trichromatic colour vision by opsin gene duplication in New World and Old World primates. Genome Research, 9(7), 629–638. https://doi.org/10.1101/gr.9.7.629
Gazit, I., & Terkel, J. (2003). Domination of olfaction over vision in explosives detection by dogs. Applied Animal Behaviour Science, 82(1), 65–73. https://doi.org/10.1016/S0168-1591(03)00051-0
Horowitz, A. (2009). Inside of a dog: What dogs see, smell, and know. Scribner.
Kasparson, A. A., Badridze, J., & Maximov, V. V. (2013). Colour cues proved to be more informative for dogs than brightness. Proceedings of the Royal Society B, 280(1766), 20131356. https://doi.org/10.1098/rspb.2013.1356
Peitsch, D., Fietz, A., Hertel, H., de Souza, J., Ventura, D. F., & Menzel, R. (1992). The spectral input systems of hymenopteran insects and their receptor-based colour vision. Journal of Comparative Physiology A, 170, 23–40. https://doi.org/10.1007/BF00190398
_________________
Featured image: Since dogs find it difficult to distinguish between certain reds and greens (like some humans do), we should choose toys and training aids in other colors (photo by Oleghz). Other illustrations from Dr. Cynthia Cook of Veterinary Vision Inc.
I dedicate this short reflection to my students—and, by extension, to all students worldwide.
“Live as if you were to die tomorrow. Learn as if you were to live forever.” ~Mahatma Gandhi
“Live as if you were to die tomorrow. Learn as if you were to live forever” is widely attributed to Mahatma Gandhi. The exact wording is uncertain, yet the sentiment is faithful to his views. Rajmohan Gandhi, in The Good Boatman: A Portrait of Gandhi (1995), summarises his grandfather’s position as “[…] a man should live thinking he might die tomorrow but learn as if he would live forever.” Rajmohan Gandhi, incidentally, is a professor at the University of Illinois at Urbana–Champaign, with whom we established an excellent student exchange in the early 2000s.
The idea itself is far older. Isidore of Seville (c. 560–636 CE): expressed a similar exhortation in the Etymologiae: “Study as if you were to live forever; live as if you were to die tomorrow.”1 Comparable formulations appear in Islamic tradition—for example, in a Hadith often rendered as: “Live for your afterlife as if you will die tomorrow, and live for this life as if you will live forever.” Although popular, this version is not found verbatim in the canonical collections.2
Some scholars have noted parallels in the writings of Desiderius Erasmus (1466 – 1536), who likewise encouraged a readiness for death coupled with the lifelong pursuit of learning, though no primary source confirms the wording frequently attributed to him.3
Jiddu Krishnamurti noted that “the whole of life … is a process of learning,” underscoring that education does not end with formal schooling but accompanies us until death. Seneca argued that time must be used wisely so that life does not slip away unexamined—the core of living fully in the present. Socrates famously declared that “the unexamined life is not worth living,” connecting philosophical inquiry with purposeful existence.
The message is timeless. Continue your quest for knowledge. Do not postpone learning; embrace it today rather than tomorrow, for even your smallest discovery joins the shared store of human knowledge. At times, it may seem no more than a single drop—but then, the vast oceans themselves are born of such drops.
Notes
The maxim “Disce tamquam semper victurus; vive tamquam cras moriturus” has long been attributed to Isidore of Seville and appears in standard editions of the Etymologiae. Chapter numbering may vary slightly by edition, but Book III contains the traditional formulation. ↩︎
The popular saying “Live for your afterlife as if you will die tomorrow, and live for this life as if you will live forever” is not found verbatim in the canonical Hadith collections. It appears in later moralistic literature and is classified by scholars as non-authentic (apocryphal). ↩︎
The widely circulated maxim “Live as if you were to die tomorrow; study as if you were to live forever” does not appear in Erasmus’s authenticated works. Modern quotation collections repeat the attribution without citing an original source, and Erasmus scholars consider it a later invention reflecting themes he discussed but never expressed in this form. ↩︎
References
Gandhi attribution Gandhi, R. (1995). The good boatman: A portrait of Gandhi. Viking Penguin. ISBN 9780670856150. — Paraphrase on p. 154.
Isidore of Seville Isidorus Hispalensis. (2006). Etymologiae (W. M. Lindsay, Ed.; reprint ed.). Oxford University Press. ISBN 9780199266941. — Book III, Ch. 24.
Hadith attribution (non-canonical) al-Sakhāwī, M. A. (1996). Al-Maqāṣid al-ḥasana fī bayān kathīr min al-aḥādīth al-mushtahira ʿalā al-alsina. Dār al-Kutub al-ʿIlmiyya. ISBN 9782745122486.
Erasmus attribution (misattribution) Rummel, E. (Ed.). (2004). The Erasmus reader. University of Toronto Press. ISBN 9780802085841.
Krishnamurti Krishnamurti, J. (1981). Life Ahead: On Learning and the Search for Meaning. Harper & Row.
Seneca Seneca. (2010). On the shortness of life (C. D. N. Costa, Trans.). Penguin Classics. (Original work published ca. 49 CE.)
Socrates (via Plato) Plato. (2002). Apology (G. M. A. Grube & J. M. Cooper, Trans.). In J. M. Cooper (Ed.), Plato: Complete works (pp. 17–36). Hackett Publishing.
Featured image: Even the great oceans are made of many tiny drops (photo by Nick Grabowski).
“Do dogs understand what we say?” is one of the most frequent questions people ask me.
My answer is, “yes and no. They do, and they don’t. It all comes down to what you mean by understanding.”
Dogs do not understand English or any other human-created language. They do understand words (or rather, sounds) in any language. After hearing the sound “sit” many times, the dog associates it with a particular behavior and with some consequences, and will end up sitting more often than not when it hears that sound. It all depends on the consequences and on the competing stimuli at that precise moment. If the dog has something better to do, offering more attractive consequences, or the consequences for not sitting are not that unpleasant, then it won’t sit. In that respect, it is exactly like us: “I hear perfectly well what you are saying, I just don’t want to do it.” It is all a question of costs and benefits, as we say in ethology.
Dogs do not understand sentences. Most dogs get excited and run to the door when we say, “Let’s go for a walk.” That does not prove the dog understands the sentence; it only shows that it associates one sound in the sentence—probably the word walk—with one particular behavior. If we say, “Banana ping-pong walk,” we will very likely get the same response.
Tone matters. We don’t need any experiments to verify that. Observing casual dog owners provides us with all the necessary evidence. “Don’t do that, sweetie, we don’t like that at all,” with a gentle voice, is no way to prevent a dog from doing whatever it is doing. Better be quiet if so, because all we say in that tone will only reinforce the behavior we don’t want. Curious, isn’t it, how things can work just the opposite of what we intend?
There is a universal language comprising terms that all animals understand, such as peace, danger, companionship, fear, safety, and mutuality. Partnerships exist between animals across species (photo by unknown).
If you want your dog to keep on doing what it is doing, you’d better say something in a mellow tone. It does not matter what you say, but it will be more efficient if you always use the same word (read sound). Personally, my favorite is dygtig (Danish for clever). It has a good doggy sound, gives me a friendly, doggy face, and I can modulate it for the occasion, e.g., make it long, short, etc.
If you don’t want your dog to do something, you’d better say it in a serious tone (I said serious, not yelling). I use “Stop” or “Phooey” in an assertive tone, and that does the trick (usually). I never use “No” for this purpose. “No” conveys important information, i.e., “What you’re doing is not adequate, try something else.” Of course, you don’t need to do as I do. You do what works for you, and I do what works for me.
Body language is essential, and even more decisive in our dogs’ behavior than sounds and tones. If you doubt it, watch my movie “Animal Training My Way.” I barely talk to the dog, and we understand one another perfectly well. Self-confident body language will induce your dog to follow your instructions more readily. Insecure body language will either make your dog nervous or alert it to take control of the situation since you seem to be in no position to do anything about it.
Does it help to try to speak dog language, even with an awful accent? Yes, definitely. Dogs respond well to our yawning, champing (chomping), licking our lips, squeezing our eyes shut, pouty mouth, the canine muzzle grasp, and many other signals. You need to be a keen observer and practice, and to be completely uninhibited and unconcerned about others laughing at you. I like doing it, and I get excellent results. Then again, I speak nine languages (doguese, catese, and horsish not counted), some with a poor accent—and I do get rewarded for my effort. It works for me, but again, you do what works best for you.
Do dogs create relationships with us like they do with other dogs? Not exactly, but does it matter? Dogs are uncomplicated. When they live with other animals, including humans, they adapt (as do many other animals). They don’t regard us as dogs, and I believe they don’t even speculate about that. They communicate with us in their language, and they seem to appreciate it when we answer them in something that resembles their language. There’s nothing special about that. It works for and with most animals (if not all). You respect their ways, and you get some results—you don’t, and you get different results.
It’s all a question of communication. When I’m diving with rookie students, their way of moving around and gesticulating far too much attracts the attention of the local fauna. When I’m there with one of my usual diving buddies (we always dive in buddy pairs), they don’t even seem to notice us. The body language of the rookie signals “alarm,” “intruder”—and ours, more experienced as we are, signals “all is good.”
It’s really that simple. I still can’t grasp how anyone can argue that meeting the other party halfway is pointless. The usual defense is that dogs are dogs and humans are humans—a remarkable justification that flies in the face of everything we know about interspecies communication—hence, my commitment to “knowledge to everyone everywhere.”
All I can say to you is that it works well for me. With all my human inadequacies—and within certain limits—when in Rome, I do as the Romans do; when underwater, I do as the fish do; and when I’m with a dog, I do as dogs do. You don’t have to, of course. Yet, I tell you, every time we manage to bridge that gap, even for an instant, we glimpse something larger than ourselves—the rudiments of a language that might well be universal.
____________
Featured image: Dogs communicate with us in the ways of their species, and they seem to appreciate it when we answer them in something that resembles their language (photo by Lisa Jernigan Bain).
This is a beautiful recording of a lovely moment. What strikes me most in this clip is the peace emanating from both the little boy and the dog. It is but an elusive instant in the infinite history of time, but, for all they care, the world could be in flames. That one moment they share, nothing can take from them, it is all they have there and then. It will never be undone, it will never be any different, frozen as it is for all eternity. They are what they are, and they are no different. Peace comes not from striving and desiring, but from being—no conditions, no expectations, no questioning the past or querying the future. Life is what it is, and any relationship is unique because it involves unique individuals and unique conditions.
The magic of life lies not in living against, but in living with.
__________
As much as I would like to credit the author of this clip, unfortunately, his or her name remains unknown to me. Thanks for allowing us to share this beautiful, private moment.
__________
PS— At 1730 hrs GMT, 10 hours after I published my blog, I received a message from my Facebook friend Joeson Hsu in Taiwan with the information I had missed. Thanks, Joeson. The author of this movie is Ana, the mother of Herman, the little boy, and the dog is Himalaya. Thank you so much, Ana, for sharing with us. Indeed, communication is a will, not a question of language or species, and a relationship is a natural thing.
__________
PPS from October 27, 2025—Meanwhile, Juan Cardoza uploaded the clip with all the information I had missed when I first published my blog on May 12, 2014, and this is the link I have now embedded in this blog.
Your comments are welcome. Please feel free to leave a reply.