Jul 14 2026

The Altruism Paradox Revisited: Plastic Phenotypes and the Ecology of Social Strategies

Roger Abrantes

*Two honeybees (Apis mellifera) performing trophallaxis — the mouth-to-mouth transfer of food, a classic example of altruistic behaviour in honey bees (photo from Wikipedia).*

Abstract

The persistence of altruistic behaviour has long presented a challenge for evolutionary theory because behaviours that reduce an individual’s own fitness while benefiting others appear vulnerable to replacement by selfish alternatives. Classical explanations, including kin selection, reciprocal altruism, group selection, and gene-centred approaches, have generally addressed this problem by assuming competition between genetic determinants of altruistic and selfish behaviour.

Following Yakubu’s (2013) critique of this assumption, this paper explores the possibility that altruistic and selfish behaviours are not expressions of competing alleles but rather plastic phenotypic expressions of the same genotype, their deployment depending upon ecological, developmental, and social conditions. Evidence from social insects and vertebrates is reviewed in support of this interpretation.

However, replacing competing genetic determinants with phenotypic plasticity does not entirely eliminate the evolutionary problem. It merely shifts it from the genetic to the ecological level. If altruism and selfishness are alternative expressions of the same genotype, evolutionary theory must still explain the conditions under which one behavioural strategy is favoured over another, and why both persist within populations.

The paper argues that this question belongs to the domain of evolutionarily stable strategies and frequency-dependent selection. In this view, phenotypic plasticity and ESS theory are not competing explanations but complementary ones: the former may explain the coexistence of altruistic and selfish tendencies within individuals, whereas the latter may explain their persistence within populations.

Keywords: altruism; phenotypic plasticity; evolutionarily stable strategy (ESS); behavioural ecology; phenotypic gambit; frequency-dependent selection; evolutionary game theory; ASP model

1. Introduction

Altruism refers to unselfishness, selflessness, or the regard for others as a guiding principle, in contrast to egoism or selfishness. Although the term is commonly used in everyday language, its meaning and implications are complex and have long posed significant questions for biologists, philosophers, and other scientists. Defining altruism is not as straightforward as it may initially appear. Furthermore, constructing a model in which such behaviour is both evolutionarily plausible and consistent with evolutionary theory remains a considerable challenge.

1.1. Etymology

Altruism derives from the French autrui, meaning “other people.” Autrui developed from the Old French autre = other, which itself comes from Latin alter = other (Oxford English Dictionary). The French autrui gave rise to altruisme, from which the English altruism was subsequently derived.

The English term has been in use since the middle of the nineteenth century. Its adoption was facilitated by the writings of Auguste Comte, the founder of positivism, who introduced altruisme as an antonym of egoism in his Système de politique positive (Comte, 1851–1854).

2. Definitions and Problems

In everyday language, altruism denotes unselfishness, selflessness, or concern for the welfare of others. Although these definitions appear intuitive, they become problematic when applied to biology. Before examining evolutionary explanations, it is therefore necessary to distinguish among three concepts that are frequently conflated.

First, ordinary-language altruism refers simply to behaviour intended to benefit another individual, irrespective of its underlying causes or consequences. This is the meaning most people have in mind when they describe an action as altruistic.

Second, psychological altruism concerns motivation. An action is psychologically altruistic only if it is performed solely for the benefit of another individual, without any expectation of reward or personal satisfaction. Whether such motivation exists has long been debated by philosophers and psychologists. Proponents of psychological egoism argue that even apparently selfless acts ultimately benefit the actor, for example through personal satisfaction or social approval (Slote, 1964).

Third, biological altruism is defined independently of motivation. In evolutionary biology, behaviour is considered altruistic when it increases another individual’s fitness at a cost to the actor’s own fitness. Whether such behaviour ultimately increases the actor’s inclusive fitness or is favoured through indirect mechanisms is a separate question.

Unless otherwise specified, the term altruism in this paper refers to biological altruism, since the evolutionary problem concerns fitness rather than intention.

This distinction allows us to identify many behaviours that may reasonably be interpreted as altruistic.

Parental care provides perhaps the clearest example. Parents invest considerable time and energy in feeding, protecting, and raising their offspring while incurring immediate energetic and reproductive costs. Female domestic dogs (Canis lupus familiaris), for example, nurse their pups, regurgitate food for them, defend them against predators, and guide their behavioural development over many weeks (Fox, 1972).

Among social canids, care extends beyond the parents. In golden jackals (Canis aureus), older offspring—usually female yearlings—frequently assist in rearing younger siblings by guarding, provisioning, and protecting them despite receiving no immediate reproductive benefit (Rogers & Kaplan, 2003).

However, apparent altruism does not necessarily imply altruistic mechanisms.

A striking example comes from a widely reported observation in the Samburu National Reserve, where a lioness repeatedly adopted an oryx calf rather than killing it (David Sheldrick Wildlife Trust, 2008). Although this behaviour appears extraordinarily altruistic, it is more plausibly explained as an expression of maternal behaviour triggered under unusual hormonal conditions than as an adaptation specifically favouring unrelated individuals.

Behaviour that appears altruistic may therefore arise from mechanisms unrelated to altruism itself. These examples illustrate an important methodological principle: behaviour should not be classified as altruistic solely on the basis of its apparent outcome. Similar behavioural patterns may arise from different biological mechanisms.

These difficulties have accompanied the study of altruism since the beginnings of evolutionary theory.

Darwin recognised the problem explicitly in The Descent of Man:

“It is extremely doubtful whether the offspring of the more sympathetic and benevolent parents, or of those who were the most faithful to their comrades, would be reared in greater numbers than the children of selfish and treacherous parents belonging to the same tribe. He who was ready to sacrifice his life, as many a savage has been, rather than betray his comrades, would often leave no offspring to inherit his noble nature.” (Darwin, 1871, p. 163)

3. The Altruism Paradox

In classical formulations of the altruism problem, evolutionary biologists typically assume the existence of a gene that causes its bearer to behave altruistically toward others, while individuals lacking that gene behave selfishly. As Samir Okasha writes, “the altruists will be at a fitness disadvantage, so we should expect the altruistic gene to be eliminated from the population” (Okasha, 2009).

In other words, how can an altruistic gene, coding for a phenotype that incurs fitness costs, survive competition against a selfish gene coding for a phenotype with higher direct fitness? This is the altruism paradox.

As we shall see, there may be a way around this difficulty. Before exploring that possibility, however, we must first review the traditional evolutionary explanations. Whether such a way around removes the paradox entirely remains an open question.

4. Classical Evolutionary Explanations

4.1 Group selection

Group selection is a proposed mechanism of evolution in which natural selection operates at the level of the group rather than exclusively at the level of the individual (Wynne-Edwards, 1962; Lorenz, 1963).

Darwin, in The Descent of Man (1871), attempted to explain the evolution of human altruism as a selection process acting at the group level:

“When two tribes of primeval man, living in the same country, came into competition, if (other things being equal) the one tribe included a great number of courageous, sympathetic and faithful members, who were always ready to warn each other of danger, to aid and defend each other, this tribe would succeed better and conquer the other.” (Darwin, 1871, p. 162)

Darwin’s explanation may strike the modern reader as somewhat non-Darwinian, illustrating the magnitude of the problem he was attempting to solve.

A winner has no evolutionary impact per se unless it possesses greater Darwinian or inclusive fitness, that is, a greater genetic contribution to the next generation’s gene pool relative to the population average. An altruist may prevail in a particular contest, but if its Darwinian and inclusive fitness are nil, altruism ends with it. If altruism survives even though altruistic individuals perish without leaving offspring, then the expression survival of the fittest cannot be interpreted literally, for the fittest are ultimately those who leave the greatest number of copies of themselves in successive generations.

This issue has concerned many researchers since Darwin, among them Hamilton (1964). As Wilson & Wilson (2007, p. 329-330) write, “something more than natural selection within single groups is required to explain how altruism and other group-advantageous traits evolve by natural selection.”

For natural selection to favour altruism in a broader context, the within-group disadvantage of altruists must be offset by the between-group advantage enjoyed by groups containing altruists (Taylor & Nowak, 2007). As Nowak (2012) and Allen et al. (2012) observe, “cooperation is always vulnerable to exploitation by defectors; hence, the evolution of cooperation requires specific mechanisms that allow natural selection to favour cooperation over defection.”

For group selection to be viable, variation between groups must exceed variation within groups. Since selection acts upon phenotypes, competition and selection may occur at multiple organisational levels. Accordingly, Wilson (2015, p. 145) argues that:

“At all scales, there must be mechanisms that coordinate the right kinds of action and prevent disruptive forms of self-serving behaviour at lower levels of social organisation.”

He summarises the issue succinctly:

“Selfishness beats altruism within groups. Altruistic groups beat selfish groups. Everything else is commentary.” (Wilson, 2015, p. 70)

As we shall see, not everyone agrees with this conclusion. Between-group selection is possible in principle, although it is generally considered weaker than selection occurring within groups. Therefore, if we are to explain behaviour performed for the good of the group, we may need to do so without relying exclusively on group selection.

Indeed, many influential explanations for the evolution of cooperative and altruistic behaviour—including kin selection, reciprocal altruism, and gene-centred selection—were originally developed as alternatives to classical group selection.

4.2 Kin Selection

To explain altruism, we must find a way for natural selection to favour altruistic genes, as proposed by the theory of kin selection (Hamilton, 1964). Kin selection is an evolutionary mechanism whereby behaviours that reduce an individual’s direct fitness may nevertheless evolve because they increase the reproductive success of genetically related individuals. It is, therefore, a form of kin altruism based on inclusive fitness. The term kin selection was introduced by Maynard Smith (1964).

Darwin had already anticipated the basic idea in The Origin of Species (1859), arguing that a selective advantage accruing to “the same stock” could allow a trait to evolve even if it proved costly to particular individuals (Darwin, 1859). Several decades later, Ronald A. Fisher and J. B. S. Haldane formalised the mathematical foundations of kin selection (Fisher, 1930; Haldane, 1932). According to Maynard Smith, Haldane famously summarised the implications of his calculations by remarking that he would be prepared to lay down his life for “two brothers or eight cousins” (Maynard Smith, 1976).

Hamilton’s inclusive fitness rule states that kin selection will increase the frequency of an altruistic trait when the genetic relatedness between recipient and donor multiplied by the reproductive benefit to the recipient exceeds the reproductive cost to the donor:

rB > C

where:

r = genetic relatedness of the recipient to the donor,
B = reproductive benefit gained by the recipient,
C = reproductive cost to the donor.

Thus, kin selection may be regarded as a particular consequence of gene selection. According to the theory, the degree to which altruistic behaviour should be extended toward others depends on their coefficient of relationship to the actor. There are two principal ways in which this may occur: through kin recognition and through association with nearby relatives (Hamilton, 1964).

Kin selection is not the same as group selection. In kin selection, genes spread because they benefit genetically related individuals carrying copies of those genes. In group selection, by contrast, a genetic trait becomes widespread because it benefits the group as an entity.

4.3 Reciprocal Altruism

Reciprocal altruism is any behaviour in which a donor incurs an immediate fitness cost while increasing a recipient’s fitness, with the expectation of future reciprocation (Trivers, 1971). The reduction in fitness is therefore temporary rather than permanent. The mechanism resembles the tit-for-tat strategy familiar from game theory (Rapoport, 1966).

Reciprocal altruism, often regarded as an instance of the prisoner’s dilemma (Mérő, 1998), is evolutionarily viable if the probability of repeated interactions between individuals is sufficiently high or if interactions continue over a sufficiently long period (Trivers, 1971).

According to Trivers (1971), reciprocal altruism can evolve only if several conditions are fulfilled:

individuals must have repeated opportunities for reciprocation;
they must be able to recognise one another as individuals;
they must remember previous interactions and obligations; and
they must be motivated to reciprocate.

At first sight, such a system appears evolutionarily unstable, since selfish individuals could exploit altruists without reciprocating. Evolutionary theory has therefore sought mechanisms capable of maintaining cooperation in the face of cheating.

One proposed mechanism draws parallels between altruistic behaviour and exaggerated sexual ornaments. Both are costly in terms of fitness and readily observable. Consequently, both may function as honest signals maintained through the handicap principle. According to this principle, honest communication is costly to the signaler and can therefore be afforded only by high-quality individuals. Receivers can rely on the signal precisely because lower-quality individuals cannot sustain such costs (Fisher, 1930; Zahavi, 1975, 1977).

Another possible stabilising factor is the homogeneity norm. Changes in phenotype and function resulting from non-silent mutations often stand out within a population. Sexual individuals may therefore preferentially select mates exhibiting fewer unusual or minority characteristics. Given sufficient time, populations may consequently evolve relatively homogeneous morphological and behavioural repertoires. In a similar way, the behavioural repertoire of a population will become evolutionarily stable once it has developed to be as homogeneous as is the rule in most species. This includes any altruistic and cooperative features (Koeslag, 1997).

4.4 Gene-Centred Selection (The Selfish Gene)

When parents sacrifice themselves for their offspring, they may still benefit genetically because the coefficient of relatedness between parent and offspring is 0.5. From this perspective, apparently altruistic behaviour can be interpreted as genetically selfish. This is essentially Haldane’s extrapolation from kin selection.

For example, in wolves (Canis lupus lupus), it may pay for a parent to sacrifice its life to save two offspring because this, on average, preserves the equivalent of one complete copy of its own genome (Mech & Boitani, 2003). The calculation becomes more complicated, however, if we consider the offspring’s probability of survival in the absence of parental care. Sacrificing their lives to save one-week-old cubs would often be a poor evolutionary investment because the cubs are unlikely to survive without parental support. Under such circumstances, the better strategy may be for both parents to survive and retain the possibility of producing future offspring.

This model helps explain why individuals generally sacrifice more for their own offspring than for the offspring of relatives or unrelated individuals (Abrantes, 2015). What advocates of the selfish-gene approach emphasise, in agreement with kin selection and inclusive fitness theory, is that the primary unit of selection is the gene (Williams, 1966; Dawkins, 1976/1989).

In the gene-centred view, the gene is the unit of replication, whereas the organism is the vehicle through which genes interact with the environment and upon which selection acts directly. As Dawkins writes, “Natural selection favours some genes rather than others not because of the nature of the genes themselves, but because of their consequences—their phenotypic effects” (Dawkins, 1976/1989).

Because genes are selfish, they tend to promote selfish behaviour in the organisms they produce. Selfish, however, is used here in a strictly evolutionary sense, referring to the tendency of genes to maximise their own replication. Dawkins writes:

“Gene selfishness will usually give rise to selfishness in individual behaviour. However, there are special circumstances in which a gene can achieve its own selfish goals best by fostering a limited form of altruism at the level of individual animals.” (Dawkins, 1976/1989)

The selfish-gene model provides a powerful explanation for kin selection and eusociality. An organism may act against its immediate individual interests because, by supporting the reproduction of related individuals, it assists copies of its genes—or genes with similar phenotypic effects—in other bodies to replicate. Thus, selfish genes may produce unselfish organisms.

The survival of each gene depends on the survival of many others. Consequently, selfish genes are not necessarily uncooperative genes. To be successful, a gene must cooperate with the other genes that share its vehicle. Genes cooperate in building organisms because they share the same route into future generations. An organism is therefore a vehicle constructed by a cooperative of genes.

Vehicles are important, but replicators are essential. Darwinian natural selection remains conceivable without vehicles but not without replicators. Indeed, when life first emerged, there may have been replicators without vehicles.

From the selfish-gene perspective, a group is not a replicator because there is no equivalent of a gene pool at the group level. Nor is a group a vehicle, since the genes within a group do not necessarily share a common route into future generations.

Accordingly, selection acting at the group level is generally expected to be weaker than selection acting at the individual or gene level. Although simple forms of group selection have received little empirical support, more sophisticated multilevel formulations have proven useful in particular cases (Wilson & Wilson, 2007).

The debate remains unresolved. Wilson argues that, although the selfish-gene approach has become widely accepted, he, Martin Nowak, and Corina Tarnita demonstrated that inclusive fitness theory is “both mathematically and biologically incorrect” and that group selection provides a more realistic model of social evolution (Nowak et al., 2010; Wilson, 2012).

Dawkins rejects the replacement of kin selection with group selection. He does not deny that group selection may occur, but argues that even in those cases where it is applicable, it is cumbersome, time-consuming, and obscures what would otherwise be a straightforward evolutionary explanation.

Both Dawkins and Wilson agree that successful genes tend to prosper and replicate, and that group living may provide important evolutionary advantages under certain circumstances. Their disagreement concerns primarily the level at which selection is most profitably analysed.

Perhaps a consensus may eventually emerge through simulations that employ narrower, more precise definitions of groups and levels of selection, as illustrated by the cellular group-selection simulations of Markvoort et al. (2014).

5. Reformulating the Altruism Paradox—A Workaround

The classical formulation of the altruism paradox assumes competition between altruistic and selfish genetic strategies.

Perhaps this confirms an old philosophical intuition: an answer cannot be better than the question it addresses. What if we are asking the wrong question—and thereby creating the paradox ourselves?

The assumption that a distinct altruistic allele risks being overrun by a distinct selfish allele lies at the heart of all the models reviewed above.¹

As Dawkins puts it, “cheat genes” are spreading through the population while “sucker genes” are driven to extinction (Dawkins, 1976/1989, p. 184). E. O. Wilson formulates the problem in similar terms: “How might such a behaviour evolve if the genes promoting it are at such a disadvantage in competition with genes that oppose it?” (Wilson, 2005, p. 159).

But suppose there are no two genotypes coding for the competing phenotypes of selfishness and altruism. Suppose instead that both phenotypes are alternative expressions of a single genotype, carrying both possibilities—not two competing alleles but one genotype with multiple behavioural options.

That is exactly what Yakubu (2013) does with his Altruism–Selfishness Plasticity (ASP) model. However, even if this reformulation succeeds at the genetic level, the question of why behavioural variation persists within populations remains, as we shall see.

For the sake of argument, let us therefore set aside, for a moment, the prevailing Altruism–Selfishness Allelomorphism (ASA) framework and explore what the ASP approach may contribute to the debate.²

6. Altruistic and Selfish Phenotypes as Plastic Expressions of a Single Genotype

Indeed, to consider altruistic and selfish phenotypes as plastic expressions of a single genotype rather than products of competing alleles (Yakubu, 2013) may provide a way around the altruism paradox.

Phenotypic plasticity is the property of a genotype to produce different phenotypes in response to different environmental conditions (Pigliucci, 2001). There is nothing unusual about such a process. Biologists have long recognised that a single genotype may give rise to multiple phenotypic outcomes depending on developmental, ecological, or social circumstances.

Originally, the term referred mainly to developmental modifications of morphological characters. Today, however, it encompasses virtually all phenotypic responses to environmental conditions, including morphological, physiological, and behavioural changes, as well as processes such as acclimatisation and learning (Kelly et al., 2012).

Several lines of evidence are consistent with the ASP model. It is entirely plausible that no distinct genotype codes either the altruistic or the selfish phenotype and that both are instead plastic expressions of the same genotype, deployed under different environmental circumstances.

If this is so, then the two phenotypes are not competitors at all. The evolutionary problem would then concern not the survival of competing genes but the circumstances governing alternative behavioural expressions. To borrow Dawkins’ metaphor, the same genotype may possess the plasticity to express itself either as a “cheater” or as a “sucker”, depending on ecological and social contingencies.

Yakubu (2013) provides substantial evidence consistent with such a reformulation of the altruism paradox. Let us review some of it.

7. Empirical Support for ASP

7.1 The Social Hymenoptera

The social Hymenoptera are favourites of evolutionary biologists because they are both highly social and haplodiploid. A honeybee (Apis mellifera) colony comprises three castes: a queen, a few hundred males (drones), and thousands of nonreproductive females (workers). The queen’s role is reproduction, while the drones’ role is to mate with queens. The workers, by contrast, maintain the colony, forage, care for the brood, and defend the nest against intruders. The reproductive queen represents the selfish phenotype, whereas the sterile workers are perhaps the closest approximation to pure altruism found in nature.

We may therefore ask whether distinct alleles for altruism and selfishness exist in eusocial populations, as classical formulations of the altruism problem implicitly assume. The evidence suggests otherwise.

Whether a larva develops into a queen or a worker depends initially on where the egg is laid and subsequently on the nutrition the larva receives (Winston, 1987). Larvae reared in queen cells are fed royal jelly (Prete, 1990), whereas larvae reared in worker cells receive worker food. Remarkably, the former become queens while the latter become workers. The eggs and young larvae are therefore developmentally totipotent—or perhaps multipotent, according to this author’s interpretation (Winston, 1987).

The honeybee provides even stronger evidence for phenotypic plasticity. Eggs and larvae transferred from queen cells to worker cells, or vice versa, will develop according to the nutritional and environmental conditions they experience, provided the transfer occurs sufficiently early in development (Winston, 1987).

Thus, the queen-worker distinction does not arise from distinct genotypes. Whether an individual becomes a selfish reproducer or an altruistic worker depends primarily upon environmental cues.

Caste differentiation occurs through epigenetic mechanisms whereby non-heritable factors influence gene expression. Both queens and workers arise from the same genome, while royal jelly acts as the principal environmental trigger. The genes encoding the major royal jelly proteins constitute one of the clearest examples of gene families acquiring novel functions during the evolution of sociality (Albert et al., 1999).

Researchers have sequenced the honeybee genome (Honeybee Genome Sequencing Consortium, 2006), and several genes whose differential expression contributes to queen and worker development are now known. Evans and Wheeler (1999) identified transcripts of seven genes that are differentially expressed in queen-destined and worker-destined larvae during critical stages of development. These findings provide a particularly striking example of phenotypic plasticity mediated by differential gene expression.

7.2 Social Environmental Circumstances

Let us now consider non-eusocial social organisms. Trivers explained the altruistic behaviour of vampire bats within the framework of reciprocal altruism (Trivers, 1971). However, the same observations can also be interpreted without postulating distinct altruistic and selfish genotypes.

Hungry vampire bats frequently solicit food from better-fed individuals. Sometimes the recipient receives regurgitated blood; sometimes the donor refuses to share.

Trivers was undoubtedly correct regarding the reciprocal nature of this behaviour. Whether an individual shares blood depends largely on whether the recipient has shared previously or is expected to reciprocate in the future (Wilkinson, 1984). The same individual may therefore behave altruistically under some circumstances and selfishly under others. The remaining question is why populations nevertheless differ so consistently in the relative frequencies of these behaviours.

This observation is entirely compatible with reciprocal altruism (Yakubu, 2013). However, because genotypes do not change from one day to the next, it appears more parsimonious to interpret these behavioural alternatives as plastic expressions of the same genotype.

Similar patterns occur in other social species. Adult male olive baboons (Papio anubis) assist group members based on prior interactions and expectations of future cooperation (Packer, 1977).

The phenomenon of efficient coercion provides another example. In ten social insect species studied by Wenseleers and Ratnieks (2006), social sanctions maintained altruistic behaviour under circumstances in which selfish behaviour would otherwise have been expected.

Altruistic individuals may also become selfish when ecological and social conditions change. In white-fronted bee-eaters (Merops bullockoides), young males often become helpers because older males prevent them from establishing their own nests. Some later become breeders themselves and cease helping behaviour (Emlen & Wrege, 1992).

Similarly, lower-ranking female meerkats may behave cooperatively while subordinate but become reproductively competitive upon attaining higher rank, sometimes killing the offspring of subordinate females (Young & Clutton-Brock, 2006).

Social organisms may therefore assume subordinate roles not because they possess altruistic genes, but because those roles constitute the best available strategy under prevailing conditions (Gadagkar, 1997).

Taken together, these examples are consistent with the hypothesis that altruism and selfishness represent alternative phenotypic expressions available to the same genotype. Individuals behave altruistically under some ecological and social conditions and selfishly under others.³ No evidence has yet demonstrated that only certain individuals are capable of altruistic behaviour, while others are capable only of selfish behaviour, when exposed to the same circumstances (Yakubu, 2013).

8. Canines as an Illustration

Canines provide a familiar illustration of the behavioural flexibility discussed above. They readily cooperate against rivals, a widespread form of territorial behaviour among companion dogs, free-ranging dogs, and wild canids (Mech & Boitani, 2003; Fox, 1972). They also defend their household, including human and non-human members of the family group. Females invest heavily in rearing and protecting their offspring, often at considerable energetic cost to themselves (Fox, 1972).

These behaviours may reasonably be interpreted as altruistic, notwithstanding the conceptual difficulties surrounding the term discussed earlier. At the same time, the very same individuals may compete fiercely over food, mates, territory, or social status (Abrantes, 2015; Mech & Boitani, 2003).

Thus, canines appear capable of expressing both cooperative and selfish behavioural strategies according to ecological and social circumstances. A cautious conclusion would therefore be that dogs exhibit forms of reciprocal altruism and cooperation under appropriate conditions, precisely as the Altruism–Selfishness Plasticity (ASP) model predicts. The persistence of such behavioural variation, however, still requires an evolutionary explanation at the population level.

9. Beyond ASP: Plasticity, Bias, and Evolutionarily Stable Strategies

The ASP model resolves an important difficulty in the classical formulations of altruism by removing the assumption that altruistic and selfish behaviours are encoded by competing alleles. Instead, both become alternative phenotypic expressions available to the same genotype and deployed according to ecological, developmental, and social circumstances (Yakubu, 2013).

However, this reformulation does not eliminate the evolutionary problem altogether. It merely relocates it.

If altruism and selfishness are not competing genes, we must still explain why populations continue to exhibit both behavioural tendencies. Why do some individuals, groups, and even species appear consistently more cooperative, submissive, or altruistic, while others display a greater tendency toward competition, dominance, or selfishness?

Part of this variation is undoubtedly attributable to environmental influences and learning history. Experience matters. Development matters. Social circumstances matter. Yet these factors alone do not seem sufficient to explain the full extent of the observed differences.

Behavioural ecologists have long recognised that individuals differ consistently in their behavioural tendencies. Some individuals behave more boldly than others, some more aggressively, some more tolerantly, and some more cooperatively. Such differences often persist over long periods and frequently show significant heritability (Dingemanse et al., 2010).

This suggests that although there may be no genes that directly encode altruism or selfishness, there may nevertheless be genetic influences that bias the probability of expressing one behavioural option rather than another (Pigliucci, 2001; Kelly et al., 2012).

The distinction is important.

Genes need not determine whether an individual behaves altruistically or selfishly in a particular encounter. Instead, they may influence thresholds, sensitivities, reaction norms, and response probabilities. Two individuals may possess the same behavioural repertoire while differing substantially in the circumstances under which particular behaviours are expressed.

To illustrate, two wolves may both be capable of cooperation, submission, competition, and aggression. One may cooperate readily and escalate reluctantly, whereas another may escalate quickly and cooperate only under strong incentives. Both individuals possess the same behavioural options. What differs are the probabilities and thresholds governing their expression (Abrantes, 1997; Mech & Boitani, 2003).

Thus, the issue may not concern genes for altruism versus genes for selfishness, but rather genes influencing the probability that one strategy will be deployed rather than another under particular ecological and social conditions.

If this is correct, then the problem returns to the domain of evolutionary game theory and evolutionarily stable strategies (Maynard Smith, 1964; Nowak, 2012).

Natural selection is indifferent to the genetic mechanisms underlying behaviour. It operates upon phenotypes. What succeeds tends to persist; what fails tends to disappear.

In this respect, we are back to the spirit of the phenotypic gambit (Grafen, 1984). Evolution need not concern itself with the precise genetic architecture underlying behaviour. It acts upon phenotypes and behavioural strategies because, as always, what works, works.

Consequently, the persistence of altruistic and selfish tendencies within populations may depend less upon their genetic implementation than upon their costs and benefits under prevailing ecological and social conditions. Under some circumstances, selfish strategies will outperform altruistic ones and increase in frequency. However, if selfishness becomes too common, the social environment itself may deteriorate to such an extent that cooperative strategies once again become advantageous. Conversely, populations dominated by altruistic individuals may create opportunities for exploitation by selfish individuals, thereby favouring the re-emergence of selfish strategies.

The result may be a dynamic equilibrium maintained by frequency-dependent selection rather than by the persistence of competing genes (Taylor & Nowak, 2007; Allen et al., 2012).

From this perspective, the Darwinian algorithm remains entirely intact. The object of selection simply changes.

If so, the ASP model and ESS theory are not competing explanations but complementary ones. ASP may help explain why altruism and selfishness coexist within an individual’s behavioural repertoire. ESS may add a further level of explanation for why both tendencies continue to persist within populations.

The altruism paradox may therefore disappear at the genetic level while appearing to re-emerge at the ecological level, but only as a pseudo-paradox, for it is in reality simply a problem of strategy rather than heredity.

10. Conclusion

This paper has argued that the altruism paradox may arise from a shared assumption underlying the classical models: namely, that altruistic and selfish behaviours are encoded by distinct competing genetic strategies. Following Yakubu (2013), we have explored the possibility that both are better understood as plastic expressions of the same genotype, their expression determined by ecological, developmental, and social conditions.

The ASP model is an alternative explanatory framework rather than a replacement for kin selection, reciprocal altruism, group selection, or gene-centred approaches. These theories may continue to explain important aspects of social evolution even if altruistic and selfish behaviours prove to be plastic expressions of the same genotype.

At the genetic level, the altruism paradox may therefore disappear. If there are no altruistic genes competing against selfish genes, there is no longer any genetic competition to explain.

At the ecological level, however, the problem remains very much alive. Altruism exists. Selfishness exists. Both persist, and their frequencies vary enormously among individuals, groups, populations, and species.

The question therefore changes rather than disappears. Instead of asking how altruistic genes survive competition with selfish genes, we ask under what ecological and social conditions altruistic and selfish phenotypes are favoured and maintained.

That is ESS territory.

It returns the problem to the domain of evolutionarily stable strategies, frequency-dependent selection, and the Darwinian algorithm itself. ASP may explain why altruism and selfishness coexist within individuals. ESS may help explain why both continue to coexist within populations.

The implications of this perspective may extend beyond altruism. Other behaviours traditionally regarded as genetically distinct phenotypes may likewise prove to be plastic expressions of a common genotype whose expression is biased by heredity and shaped by developmental, ecological, and social conditions.

Perhaps the altruism paradox was never asking how altruistic genes survive selfish ones, but under what circumstances evolution favours one expression of life over another.

Sometimes the difficult part is not finding the answer. It is discovering which question evolution has actually been answering all along.

Footnotes

Yakubu (2013, pp. 105–106) notes that the conventional approach to altruism “implies that the altruist and non-altruist are distinguishable genetically by the possession (or lack thereof), of ‘a gene for altruism.’ In other words, we assume a genetic basis for the contrasting behaviors of altruism and selfishness.” ↩︎
Yakubu (2013, p. 106) writes: “In modeling the evolution of altruism, extant models build upon this template, in which they usually assume altruism and selfishness to be the contrasting alleles A1 and A2—the phenotypic gambit.” He refers to this framework as the Altruism Selfishness Allelomorphism (ASA) and contrasts it with the Altruism Selfishness Plasticity (ASP) model, in which altruism and selfishness are “alternative phenotypes of a single plastic genotype.” ↩︎
Discussing social organisms, Yakubu (2013, p. 112) observes that “individuals are often able to make transitions between the two phenotypes in response to changes in their social environment,” a finding he regards as inconsistent with models assuming separate altruistic and selfish genotypes. ↩︎

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Conflict of Interest Statement
The author declares that the research was conducted in the absence of any political, commercial or financial relationships that could be construed as a potential conflict of interest.

Jun 25 2026

Motivation—Where There Is a Will, There Is a Way

—A Conceptual Review in Ethology and Behavioural Science

Abstract

The concept of motivation is widely employed across ethology, behavioural science, psychology, and animal welfare research, yet it remains theoretically problematic and resistant to precise definition (Colgan, 1989; McFarland, 1989). Although often treated as a causal explanation of behaviour, motivation is better understood as an inferred construct that summarises the internal and external conditions under which goal-directed behaviour occurs (Dawkins, 1990). This review examines the historical development and theoretical uses of the term motivation within ethology and related disciplines, tracing its roots from instinct theory and drive-reduction models to contemporary regulatory and optimisation approaches. It argues that while motivation remains indispensable as a descriptive and heuristic concept, it cannot be exhaustively reduced to singular physiological or psychological mechanisms. A pluralistic and cautious approach is therefore warranted.

1. Introduction

Motivation is a term used so routinely that it is often assumed to be self-explanatory. In both scientific and everyday discourse, we speak as though its meaning were transparent and shared. Yet closer inspection reveals that motivation is conceptually ambiguous, theoretically overloaded, and frequently used as a placeholder for incomplete explanations (Colgan, 1989). The present review examines motivation not as a unitary mechanism, but as a functional and inferential construct employed to account for organised, goal-directed behaviour (McFarland, 1989).

For the purposes of this review, motivation is the functional organisation of behaviour arising from the interaction of an organism’s internal state, developmental history, and current environment such that behaviour is organised in ways that serve biologically and ecologically relevant functions.¹ Rather than invoking subjective experience, conscious intention, or specific neural mechanisms, this definition identifies the explanatory domain within which the concept of motivation is employed (Tinbergen, 1963).

Figure 1. Conceptual representation of motivation as the functional organisation of behaviour. Motivation is conceived as arising from the interaction of an organism’s internal state, developmental history, and current environment, such that behaviour is organised in ways that serve biologically and ecologically relevant functions.

2. Early Models: Instinct and Drive

2.1 Instinct-Based Explanations

Early biological accounts of behaviour relied heavily on the concept of instinct. Influenced by evolutionary theory, behaviour was understood as the expression of inherited programmes shaped by natural selection (Darwin, 1859/2009). Survival and reproduction were treated as ultimate explanatory principles, with instinct serving as the proximate mediator (Lorenz, 1950).

Classical ethology formalised this approach. Ethologists defined instincts as species-typical, unlearned behaviour patterns elicited by specific stimuli (Lorenz, 1950; Tinbergen, 1951). Motivation, within this framework, was often conceptualised as the activation of innate releasing mechanisms or the accumulation of action-specific energy (Lorenz, 1981).

While heuristically useful, instinct-based explanations proved theoretically unstable. As Tinbergen (1963) noted, any unexplained behaviour could be accommodated by postulating a new instinct, rendering the concept effectively unfalsifiable.

2.2 Drive-Reduction and Homeostasis

To address these limitations, motivation was reconceptualised as internal drives. Drive-reduction theory proposed that organisms possess physiological needs—such as hunger or thirst—that generate states of arousal, motivating behaviour aimed at restoring homeostasis (see Colgan, 1989).

Early ethologists adopted modified versions of this framework, frequently employing hydraulic or energy-accumulation metaphors to describe motivational states (Lorenz, 1981). Although these models clarified certain behavioural regularities, they struggled to account for exploratory, play, and curiosity-driven behaviours that increase, rather than reduce, stimulation (McFarland, 1989).

3. Beyond Minimisation: Optimisation and Regulation

Subsequent theoretical developments rejected the assumption that organisms invariably seek to minimise stimulation. Instead, motivation came to be understood as the regulation of activity towards optimal levels of engagement with the environment (McFarland, 1989; Colgan, 1989). These models more readily accommodate behaviours such as exploration, novelty-seeking, and information gathering, which may themselves constitute functionally adaptive components of an organism’s extended phenotype (Dawkins, 1982).

Physiological regulation theories further emphasised the role of complex neural systems in coordinating motivational processes (Salamone & Correa, 2012; Salamone, 2024). However, even these accounts stop short of offering a complete explanation. Neural correlates of motivation describe how behaviour is regulated, not why particular goals are selected or prioritised (Dawkins, 1990).

4. Motivation, Learning, and Communication

Motivation is inseparable from learning processes.² Reinforcement and punishment alter behaviour only insofar as the organism is motivated to engage with relevant stimuli (Breland & Breland, 1961; Catania, 1984). Apparent failures of conditioning often reflect motivational constraints rather than learning deficits, as demonstrated by instinctive drift (Breland & Breland, 1961) and by contemporary experimental work distinguishing learning capacity from motivational investment (Meagher et al., 2020).

Similarly, communication behaviours presuppose motivational states that render signalling functional in specific ecological and social contexts (Dawkins, 1990).

Strictly reflexive responses—such as simple Pavlovian reflexes—are sometimes cited as exceptions. Yet even here, contextual modulation and motivational gating suggest that motivation cannot be entirely excluded from explanatory accounts (Colgan, 1989).

5. Evolutionary and Sociobiological Perspectives

Sociobiology emerged as an attempt to ground motivation more firmly in evolutionary theory by integrating genetic, ecological, and behavioural data (Wilson, 1975). From this perspective, motivation reflects evolved behavioural tendencies shaped by selection pressures, rather than discrete internal drives.

While this approach clarified ultimate explanations, it did not eliminate conceptual ambiguity at the proximate level. Motivational terms often serve as shorthand for fitness-related behavioural patterns rather than as mechanistic explanations (Dawkins, 1982, 1990).

6. Motivation and Animal Welfare

In applied contexts, particularly animal welfare science, motivation is frequently invoked to justify claims about behavioural needs.³ Hughes and Duncan (1988) cautioned that the notion of ethological “need” must be used carefully, as motivational strength cannot be inferred directly from observed behaviour alone.

Within animal welfare science, motivation functions as both a descriptive and a normative concept, linking behavioural expression, welfare assessment, and ethical evaluation (Dawkins, 1990; Coria-Avila, 2022).

7. Conceptual Limits and Theoretical Caution

Despite more than a century of theoretical development, no single model of motivation has proven sufficient. Contemporary consensus recognises motivation as a necessary but incomplete explanatory construct, summarising behavioural regularities without fully specifying their causal architecture (Colgan, 1989; McFarland, 1989).

Accordingly, motivation should be treated neither as a literal internal entity nor as a purely linguistic convenience. Its value lies in its heuristic function—guiding inquiry while remaining open to revision (Tinbergen, 1963).

8. Conclusion

As an explanatory construct, motivation remains central to the study of behaviour precisely because it resists simple definition. Attempts to reduce it to instincts, drives, neural circuits, or genetic programmes have each illuminated certain aspects of behaviour while leaving others unexplained. A pluralistic and theoretically cautious approach—sensitive to both proximate and ultimate levels of analysis—remains the most defensible position at present. Rather than representing a single causal mechanism, motivation is best viewed as a conceptual framework that links physiological processes, behavioural organisation, and evolutionary function, thereby providing an indispensable explanatory framework within ethology and behavioural science.

Footnotes

In this review, motivation is treated as an explanatory and heuristic construct rather than as a directly observable or unitary causal variable. References to motivational states are therefore not intended to imply the existence of a discrete internal entity, but to summarise the functional organisation of behaviour under particular internal and external conditions. This usage follows a long-standing ethological tradition and is consistent with multilevel analyses that distinguish proximate mechanisms from ultimate explanations (Tinbergen, 1963; Colgan, 1989; McFarland, 1989). ↩︎
The present review does not treat motivation, drive, reinforcement, or learning as interchangeable explanatory terms. Motivation is used at a functional-descriptive level to characterise the conditions under which behaviour is organised and expressed, whereas reinforcement and punishment refer to empirically defined learning processes that modify behaviour contingent on its consequences. References to drives are retained only in their historical context. No claim is made that motivational constructs can be reduced to, or derived directly from, reinforcement histories or physiological variables alone (Breland & Breland, 1961; Catania, 1984; Colgan, 1989). ↩︎
References to motivation in the context of animal welfare are not intended to imply that behavioural expression provides a direct or exhaustive measure of welfare or subjective experience. Motivational strength, preference, and welfare value are analytically distinct and may diverge under specific ecological, developmental, or experimental conditions. Motivational explanations are therefore used descriptively and comparatively, not as normative claims about welfare adequacy or ethical sufficiency (Hughes & Duncan, 1988; Dawkins, 1990; Coria-Avila, 2022). ↩︎

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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.

Dec 27 2025

Bonding and Attachment in Companion Animals: A Functional–Ethological Analysis

—A Comparative Analysis of Dogs and Horses

Abstract

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 behavioural 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 behaviour 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 behaviour, 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. ¹ ²

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 emphasised 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 specialised 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, helps regulate 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 behaviour, stress responsiveness, and affiliative tendencies (Bowlby, 1982; Carter, 1998).⁴ ⁵

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 behaviour, 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 behaviour, including shared parental investment, mate guarding, or cooperative resource defence, thereby increasing the likelihood that shared genetic material is successfully transmitted to subsequent generations.

Pair bonding is functionally and evolutionarily favoured 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.

Behaviours 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 specialised subset of bonding, characterised 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 emphasises that bonding encompasses a broader range of social relationships than either imprinting or attachment alone, and that neither 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 behaviour. 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	Any life stage	Multiple mechanisms: associative learning, repeated interaction, shared experience; neuroendocrine facilitation (e.g., oxytocin)	Variable; from transient to lifelong	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 behavioural 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 behavioural 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, synchronised activity, and allogrooming—a behaviour 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 behavioural 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 behaviour 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 behavioural 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 by shared activities and coordinated responses to environmental challenges than by 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, emphasise 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, recognising 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 behaviour 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 behaviours 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 behavioural 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). Recognising 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, emphasise 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 generalisation (Hamilton, 1964; Tinbergen, 1963). Bonding can be favoured 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. Recognising 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 behaviours that appear altruistic at the individual level can be favoured 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 favoured 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 behavioural patterns, without theoretical commitments concerning mental states or emotional experience. ↩︎
Bowlby explicitly characterises 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 behavioural system defined by function rather than by species-specific expression. ↩︎
Carter summarises 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 behaviours such as grooming, play, and cooperative defence 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 during 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, characterised by rapid acquisition, restricted to a sensitive developmental period, and relatively independent of reinforcement (Lorenz, 1935). Early formulations emphasised 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 behaviour 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. ↩︎

References

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Nov 24 2025

Stress Helps Learning and Bonding

Abstract

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).

Stress006 — 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).

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Sweatt, J. D. (2013). The emerging field of neuroepigenetics. Neuron, 80(3), 624–632. https://doi.org/10.1016/j.neuron.2013.10.023

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

Nov 15 2025

Your Dog Understands Your Yawn

Abstract

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

Argiolas, A., & Melis, M. R. (1998). The neuropharmacology of yawning. European Journal of Pharmacology, 343, 1–16. https://doi.org/10.1016/S0014-2999(97)01538-0

Gallup, A. C. (2022). The causes and consequences of yawning in animal groups. Animal Behaviour, 187, 209–219. https://doi.org/10.1016/j.anbehav.2022.03.011

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

Guggisberg, A. G., Mathis, J., Schnider, A., & Hess, C. W. (2010). Why do we yawn? Neuroscience & Biobehavioral Reviews, 34(8), 1267–1276. https://doi.org/10.1016/j.neubiorev.2010.03.008

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

Norscia, I., & Palagi, E. (2011). Yawn contagion and empathy in Homo sapiens. PLOS ONE, 6(12), e28472. https://doi.org/10.1371/journal.pone.0028472

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

Nov 6 2025

Canine Scent Detection: Reviving the Oldest Mammalian Sense

—A Sniffer Dog is a Happy Dog

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.

References

Banerjee, A., & Bhadra, A. (2022). Time–activity budget of urban-adapted free-ranging dogs. Acta Ethologica, 25(1), 15–25. https://doi.org/10.1007/s10211-021-00379-6

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

McGann, J. P. (2017). Poor human olfaction is a 19th-century myth. Science, 356(6338), eaam7263. https://doi.org/10.1126/science.aam7263

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

Shipley, M. T., & Ennis, M. (1996). Functional organization of olfactory system. Journal of Neurobiology, 30(1), 123–176. https://onlinelibrary.wiley.com/doi/10.1002/(SICI)1097-4695(199605)30:1%3C123::AID-NEU11%3E3.0.CO;2-N

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.

Sep 23 2025

The Principle of Simplicity

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…”

The principle 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.

References

McFadden, J. (2023). Razor sharp: The role of Occam’s razor in science. Annals of the New York Academy of Sciences. https://doi.org/10.1111/nyas.15086 (PMC article: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10952609/).

Walsh, D. (1979). Occam’s razor: A principle of intellectual elegance. American Philosophical Quarterly, 16(3), 241–244. https://www.jstor.org/stable/20009764.

Feb 13 2022

The Illusive Free Will

—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 Neurosci 11, 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.

Psychology Rather Than Power—for the Umpteenth Time, Reinforcers Are Not Rewards

*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.

Skinner, B. F. 1986. What is wrong with daily life in the Western world? American Psychologist, 41(5), 568-574. https://doi.org/10.1037/0003-066X.41.5.568. Retrieved Jun. 29, 2019.

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.

Apr 12 2017

The Function of Champing Behaviour: An Ethological Account in Canines

Abstract

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).

janegoodallandchimp1 — *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).

Blass, E. M. (1980). Suckling. Science, 210(4472), 729–735. https://doi.org/10.1126/science.6997992

Goodall, J. (1971). In the shadow of man. London: Collins.

Hinde, R. A. (1976). Interactions, relationships and social structure. Man, 11(1), 1–17. https://doi.org/10.2307/2800384

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.

Oct 4 2016

The Ethology of Trial and Error: Thirty-Four Illuminating Failures

“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 inhibitors² herself, 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 reinforcer reinforces—meaning it strengthens behavior (it increases; one gets more). This is intuitive and consistent with standard usage. Likewise, an inhibitor inhibits—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.

Jun 10 2014

Do Dogs See Colors? What Does It Mean for Our Training?

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).

dogcolor6-1 — *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.

Jacobs, G. H. (1993). The distribution and nature of colour vision among the mammals. Biological Reviews, 68(3), 413–471. https://doi.org/10.1111/j.1469-185X.1993.tb00738.x

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

Neitz, J., Geist, T., & Jacobs, G. H. (1989). Color vision in the dog. Visual Neuroscience, 3(2), 119–125. https://doi.org/10.1017/S0952523800004430

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

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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.

Jun 1 2014

Live as If You Were to Die Tomorrow—Learn as If You Were to Live Forever

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).

May 17 2014

Do Dogs Understand What We Say?

“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.

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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).

May 12 2014

The Little Boy and His Dog

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.

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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.

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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.

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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.

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Abstract

1. Introduction

1.1. Etymology

2. Definitions and Problems

3. The Altruism Paradox

4. Classical Evolutionary Explanations

4.1 Group selection

4.2 Kin Selection

4.3 Reciprocal Altruism

4.4 Gene-Centred Selection (The Selfish Gene)

5. Reformulating the Altruism Paradox—A Workaround

6. Altruistic and Selfish Phenotypes as Plastic Expressions of a Single Genotype

7. Empirical Support for ASP

7.1 The Social Hymenoptera

7.2 Social Environmental Circumstances

8. Canines as an Illustration

9. Beyond ASP: Plasticity, Bias, and Evolutionarily Stable Strategies

10. Conclusion

Footnotes

References

—A Conceptual Review in Ethology and Behavioural Science

Abstract

1. Introduction

2. Early Models: Instinct and Drive

2.1 Instinct-Based Explanations

2.2 Drive-Reduction and Homeostasis

3. Beyond Minimisation: Optimisation and Regulation

4. Motivation, Learning, and Communication

5. Evolutionary and Sociobiological Perspectives

6. Motivation and Animal Welfare

7. Conceptual Limits and Theoretical Caution

8. Conclusion

Footnotes

References

—A Comparative Analysis of Dogs and Horses

Abstract

Bonding—a Definition

Parent–Offspring Bonding and Attachment

Pair Bonding and Reproductive Cooperation

Social Bonding Beyond Attachment

Neurobiological Substrates of Bonding and Attachment

Bonding, Attachment, and Imprinting

Table 1—Terminological Comparison: Bonding, Attachment, and Imprinting

Bonding and Attachment in Domestic Dogs

Bonding and Attachment in Domestic Horses

Comparative Perspective: Dogs and Horses

Table 2—Imprinting, Attachment, and Bonding in Domestic Dogs and Horses

Practical Implications for Human–Animal Interaction

Conclusion

Footnotes

References

—A Sniffer Dog is a Happy Dog

References

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When terminology goes wrong

Back to the 1980s

Discovering training through ethology

Learning from the giants

Winning where no one expected us to

And just like that, I was in deeper than I thought

Notes

Reference List (First Editions in Original Language)

📦 Glossary

Reinforcer

Reward

Inhibitor

Dygtig (Danish)

Lead vs. Leash

Force-Free (in practice)

Primary Reinforcer

Conditioned Reinforcer

Abstract

Footnotes

References

Notes

References

References

Notes

References