Do Animals Show Altruism?

Do Animals Show Altruism?

Introduction |Etymology | Definition and Problems | The Altruism Paradox- Group selection—Wilson | Kin Selection—Maynard Smith | Reciprocal Altruism—Trivers | Genetic Selfishness—Dawkins | Phenotypic plasticity | An Example We All Know—Do Dogs Show Altruistic Behavior? | Conclusion and Perspectives



Altruism means unselfishness, selflessness, the regard for others as a principle (the opposite of egoism or selfishness). We all use the word in our daily language, feeling confident we know everything about altruism, its meaning, and implications, but do we? The concept poses many questions to biologists, philosophers, and other scientists. It has always done. For one, defining altruism isn’t as straightforward as the introductory paragraph above may seem to imply. Then, setting up a model where such behavior makes sense and can evolve without contradicting evolutionary theory has proven to be a real brain twister.




Altruism derives from the French autrui = other people. Autrui developed from the Old French autre = other which itself comes from Latin alter = other. [1] Under the influence of alter, the French autrui produced the altrui- of the French altruisme and the English altruism. The English term has been in use since the mid-1800s. The translation of Auguste Comte (the French founder of the philosophical school of positivism), who coined the term for an antonym of egoism, [2] contributed to spreading its adoption in English.



Definitions and Problems

What is altruism? The vague definition of the term makes us doubt whether altruistic behavior is possible at all. Note that most synonyms of ‘altruism’ are not much more enlightening—also needing some explaining. If to be altruistic, the actor must get nothing at all in return for its actions, then we may question if any act qualifies as altruistic. Self-satisfaction always seems to follow a charitable act. Therefore, according to the theory of psychological egoism, we cannot describe sharing, helping, or sacrificing as altruistic. [3]

If we define altruism as unselfish conduct in the short term, i.e., without immediate benefit, then we can find more examples of such acts. One typical case is parental behavior. Parents do plenty for their progeny with no direct advantage to themselves. Female canine mothers, including our domestic dogs, Canis lupus familiaris, spend enough time and energy looking after their pups, feeding them, and teaching them various skills. [4] They regurgitate for them and defend them if they are in danger.

In the wild social canids, pack members also support caring for youngsters without immediate benefits. In jackals, Canis aureus—a group comprising father, mother, and cubs—when a juvenile is present (almost always, a sister to the newborn), she will help rear her younger sibling. [5].

However, note that some behavior we classify as altruistic may have a different and more plausible explanation. The case (in the Samburu National Park in Kenya) of a female lion adopting an oryx calf is more likely to result from a hormonal imbalance affecting its maternal instinct, than to altruism. [6]

Last, a problem of great concern, one that had preoccupied great minds in biology, is the ‘altruism paradox.’

Darwin well knew of that. He writes, “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.” [7]



The Altruism Paradox

In explaining kin selection, evolutionary biologists imagine a gene that causes its bearer to behave altruistically toward other organisms and those without it to behave selfishly. “The altruists will be at a fitness disadvantage, so we should expect the altruistic gene to be eliminated from the population.” [8] That is, in a nutshell, the problem of altruism for the evolutionary scientist.

How can an altruistic gene, coding for the weaker altruistic phenotype, thrive in competition against a selfish one that codes for the fitter selfish phenotype? Hence, we have the altruism paradox. As we shall see, there might be a workaround this issue, but let us first review the traditional models.



Group Selection

Group selection is another proposed mechanism of evolution in which natural selection operates at the group level, instead of at the level of the individual [9][10].

Darwin, in “The Descent of Man” in 1871,[7] attempted to explain the evolution of human altruism as a selection process 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.” [7]

Darwin’s explanation seems a tad non-Darwinian, thus revealing the magnitude of the problem.

A winner has no impact on evolution, per se, unless it has a better ‘Darwinian or inclusive fitness’ (the genetic contribution of an individual to the next generation’s gene pool relative to the average for the population). An altruist may win but if its ‘Darwinian and inclusive fitness’ is nil, altruism stops with it. If altruism survives, even though donors perish leaving no progeny, the survival of the fittest is not true for the fittest are those who leave the most copies of themselves in successive generations.

This issue has bothered many since Darwin, among them, Hamilton [11]. E. O. Wilson and D. S, Wilson write: ‘‘[…] something more than natural selection within single groups is required to explain how altruism and other group-advantageous traits evolve by natural selection.” [12]

For natural selection to favor altruism in a broader scenario, the ‘within-group’ disadvantage of the altruist must be offset by the ‘between-group’ advantage of the group including altruists. [13] ‘‘Cooperation is always vulnerable to exploitation by defectors; hence, the evolution of cooperation requires specific mechanisms, which allow natural selection to favor cooperation over defection.’’ [14][15]

For group selection to be viable, we must assume that the variation between groups is larger than the variation within groups. Since selection acts upon the phenotype, competition and selection can operate at all levels. Therefore, D. S. Wilson contends that “At all scales, there must be mechanisms that coordinate the right kinds of action and prevent disruptive forms of self-serving behavior at lower levels of social organization.”[16] He summarizes, “Selfishness beats altruism within groups. Altruistic groups beat selfish groups. Everything else is commentary.”[16]

As we shall see, not everyone agrees with that. ‘Between-group’ selection is possible, in principle, although it is weak compared to any which may happen ‘within-group’. Therefore, if we are to explain ‘for the good of the group’ behavior, then we must do it without group selection.

In fact, all models for explaining how cooperative and altruistic social behavior evolve, such as kin selection, reciprocity, and the selfish gene theory developed as alternatives to group selection.



Kin Selection

To explain altruism, we must find a way of natural selection to favor altruistic genes as in the theory of kin selection.[11] Kin selection is the evolutionary strategy that supports the reproductive success of an organism’s relatives at one’s cost. It is kin altruism based on inclusive fitness. Maynard Smith used the term ‘kin selection’ for the first time in 1964.[17]

Charles Darwin discussed this strategy in “The Origin of Species,” (1859)[18] arguing that a selection benefit to “the same stock” (kin) would allow the evolution of a trait while destroying an individual. Seventy years later, Fisher and Haldane figured out the mathematics of kin selection. [19][20] According to Maynard Smith, Haldane resumed the conclusion of his calculations by saying that, “he was prepared to lay down his life for eight cousins or two brothers.” [21]

Hamilton’s inclusive fitness rule states that kin selection increases the frequency of particular genes when the genetic relatedness of a recipient to a donor multiplied by the benefit to the recipient is greater than the reproductive cost to the donor.
where r=the genetic relatedness of the recipient to the donor; B=the reproductive benefit gained by the recipient of the altruistic act; C=the reproductive cost to the donor.

Thus, kin selection is a special consequence of gene selection. The degree at which one should extend altruistic behavior toward others depends upon their coefficient of relationship. There are two ways to achieve this: (1) by kin recognition and/or (2) by living near one’s relatives (Hamilton 1964).

‘Kin selection’ is not the same as ‘group selection’ where a genetic trait may become widespread because it benefits the group as an entity.



Reciprocal Altruism

Reciprocal altruism is all behavior whereby a donor, with one act, reduces its fitness while increasing a recipient’s fitness, expecting subsequent payback. [22] Thus, the reduction is temporary. The mechanism is close to the “tit for tat” strategy in game theory.[23]

Reciprocal altruism, regarded as an instance of the prisoner’s dilemma, [24] is an evolutionary possibility if chances of meeting another reciprocal altruist are high enough, or if the game continues long enough.[22]

Reciprocal altruism, [22] is a possible way for natural selection to favor altruistic genes provided that:
(1) the individuals must have plenty of opportunities for reciprocation;
(2) they must be able to recognize each other as individuals;
(3) they must remember the obligations;
(4) and that they must be motivated to reciprocate.

Although this model seems to be evolutionarily unstable, evolutionary biologists found a way in which it would work.

There are striking parallels between altruistic behavior and exaggerated sexual ornaments. Both are costly in fitness and easy to detect, and both might be fitness signals turned evolutionarily stable by the handicap principle. The handicap principle suggests that honest communication is expensive to the signaler, therefore only affordable to special individuals. Receivers know that reliability must backup quality because lesser signallers couldn’t afford such extravagances. [25][26][27]

Then, we have the homogeneousness norm. Change in phenotype and functionality, caused by a non-silent mutation, will often stand out in a population. Thus, we can expect sexual individuals to prefer mates with the least number of unusual or minority features. As a result, and given enough time, a whole population will develop similar looks. In a similar way, the behavior repertoire of a population will become evolutionarily stable once it has developed as homogeneous as is the rule in most species. This includes any altruistic and cooperative features.[28]



Genetic Selfishness

When parents sacrifice themselves for their progeny, they are benefiting themselves since their offspring have 50% of their genes. Therefore, we can regard altruistic behavior as genetically selfish. That is Haldane’s extrapolation from ‘kin selection’.

For example, in wolves, Canis lupus lupus, [29] it pays off for each parent to sacrifice its life saving two of their progeny because this equals twice 50% of their own genes. However, such a calculation is more complicated than so if we take into account the cubs’ chances of survival without parental support. Giving their lives to save their youngsters when they are only one week old is a bad trade since they are unlikely to survive. In this instance, the best strategy is for the male and the female to protect themselves and keep the prospect of producing more offspring later.

This model explains why individuals sacrifice more for their progeny than for those of relatives or strangers, as we saw.[30] What the defenders of the selfish gene want to emphasize, besides agreeing with kin selection and inclusive fitness, is that the selection process happens at the gene level.[31][32]

In the genetic selfish model, the unit of replication is the gene, and the organism is the vehicle it uses and upon which selection acts directly. “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.”[32]

Because genes are selfish, they will promote selfish behavior in the individuals they produce. ‘Selfish’ means, in this sense, to take care of itself as the first priority. 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.”[32]

Gene selection soundly explains kin selection and eusociality. An organism acts altruistically, against its individual interests, because by supporting a related one to reproduce, genes help copies of themselves (or sequences with the same phenotypic effect) in other bodies to replicate. Thus, sometimes, ‘selfish’ actions by the genes lead to unselfish behavior by the organisms.

The survival of each gene, being replicators, depends on the survival of some others. Being a selfish gene does not imply that genes are entirely uncooperative. To be successful, a gene needs to cooperate with the other genes with which it shares a phenotype. Genes cooperate in building bodies because they all share the same exit route into the next generation. That’s their only way to survive. An organism, a body, is a vehicle for its genes, built up by a cooperative of genes.

Vehicles are important, but replicators are essential. Darwinian natural selection is still viable with no vehicles, only replicators, but not the other way around. In fact, when life began, there were probably no vehicles, only replicators.

A group is not a replicator because there is no ‘group pool’ (like there are ‘gene pools’). There is no metapopulation in which some groups are more successful than others at making replicas of themselves. And a group is not a vehicle because to qualify as one, all the genes in the same group would need to share the same exit route to the next group in the generational sequence.

Selection based upon groups is rare compared to selection on individuals, according to the selfish gene model. Researchers could not confirm simple interpretations of group selection, though more sophisticated ones proved to make accurate predictions in specific cases.[12]

E. O. Wilson writes that although the selfish-gene approach has been widely accepted “[…] Martin Nowak, Corina Tarnita, and I demonstrated that inclusive fitness theory, often called kin selection theory, is both mathematically and biologically incorrect.” He argues that group selection is a more realistic model of social evolution.[12][33[[34]

Dawkins rejects replacing ‘kin selection’ with ‘group selection.’ According to the selfish-gene model, viewing evolution as driven by the differential survival of whole groups of organisms is incoherent. He does not deny ‘group selection’. What he contends is that, even in the rare cases where it is not wrong, it is cumbersome, time-wasting, and distracting to what would otherwise be a straightforward understanding of what happens in natural selection.

Both Dawkins and Wilson agree that favorable genes are likely to prosper and replicate and that living in groups is helpful in some circumstances. The dispute arises mostly over definitions. They aim toward representing empirical facts with precision, but both use too broad definitions for ‘group,’ ‘group selection’ and ‘kin selection’. Thus, it becomes rather difficult to test their models.

We might find a consensus if we can simulate various evolutionary scenarios with narrower definitions, similar to what Markvoort et al. did for simulations of cellular group selection.[35]



A Workaround the Classic Approach

The altruism paradox, as we saw, goes down to the question: how can an altruistic gene, coding for the weaker altruistic phenotype, thrive in competition against a selfish one that codes for the fitter selfish phenotype?

Perhaps this is a confirmation that an answer cannot be better than the question it addresses. What if we are posing the wrong question and, hence, creating the paradox?

The thought of a distinct altruistic allele risking being overrun by a distinct selfish one is the basis for all the models we reviewed above.

As Dawkins writes, “‘cheat genes’ are spreading through the population while ‘sucker’ genes are driven to extinction.”[32] E. O. Wilson also summarized the problem in the same lines, “How might such a behavior evolve if the genes promoting it are at such a disadvantage in competition with genes that oppose it?” [36]

Now, suppose there are no two genotypes coding for the two competing phenotypes, selfishness and altruism. What if both phenotypes were due to one single genotype carrying both alternatives—no two distinct alleles? The Altruism Selfishness Plasticity (ASP) model suggests exactly that. Thus, let us for a moment set the prevailing Altruism Selfishness Allelomorphism (ASA) frame aside and explore what the ASP approach can add to the debate.



Altruistic and Selfish Phenotypes as Plastic Expressions of a Single Genotype

There is a way around the altruism paradox: to consider the altruistic and selfish phenotypes as plastic expressions of a single genotype. [37]

Phenotypic plasticity is the property of a genotype to produce different phenotypes in response to distinct environmental conditions.[38] There is nothing strange about that. We have known about phenotypic plasticity for quite a while. It denotes particular morphological and physiological changes in an organism—and behavior—as a response to a specific environment. Although biologists, to begin with, used the term to describe some developmental effects of a morphological character, they use it today to describe all phenotypic responses to environmental conditions, e.g., acclimatization and learning.[39]

In fact, we have plenty of data to support the ASP model. It is very plausible that no distinct genotype codes either the altruistic or the selfish phenotype, that they are rather plastic expressions of the same genotype, determined by particular environmental circumstances. And if this is true—the altruistic and selfish phenotypes being creations of a single genotype—then, they cannot be competing. To use Dawkins’ metaphor, the same genotype has the plasticity to express itself either as a ‘cheater’ or a ‘sucker’ depending on accidental contingencies.

Yakubu provides plenty of evidence in support of such a solution to the altruism paradox.[37] Let us review some of them.


The Social Hymenoptera

The social Hymenoptera are a favorite of biologists because they are highly social and haplodiploid. A honeybee, Apis mellifera, colony has three castes consisting of a queen, a few hundred males (drones), and thousands of non-reproductive females (workers). The queen’s role is to reproduce and the drone’s job is mating a queen. The workers, on the other hand, strive hard keeping the colony, foraging, and defending it from intruders. The reproductive queen is selfish, while the nonreproductive workers are perhaps the epitome of altruism.

Now, we can ask whether there are distinct alleles (genotypes) for altruism and selfishness in eusocial populations as the classical models of altruism presume. Evidence refutes that conclusively.

Whether a larva becomes a queen or a worker begins with where the egg is laid and continues with the received feeding.[40] The workers will nurse a larva in a queen cell with royal jelly.[41] On the other hand, if the larva is in a worker cell, they will feed it worker food. Amazingly enough, the former will become a queen and the latter a worker. We can say the eggs and the larva are totipotent [40] or multipotent (according to this author).

The honeybee provides us with even more staggering evidence. We can move eggs and larva from a queen cell to a worker cell, or the other way around, and if we do that within the first three days, they will develop as they are fed and as to where they reside.[40] We can thus infer that there is no genotype involved in the queen/worker distinction. Whether a bee becomes selfish (queen) or altruist (worker) depends only on environmental stimuli.

The caste differentiation happens via an epigenetic process where non-heritable factors contribute to gene expression. Queen and worker morphological forms both originate in the same genome. Royal jelly nourishment is the non-genetic determiner. The genes encoding the major royal jelly proteins present one of the clearest examples of a gene class acquiring new functions during the evolution of sociality.[42]

Researchers have sequenced the genome of the honeybee.[43] In fact, we know the particular genes in the honeybee whose differential expression results in queen and workers. Evans and Wheeler identified and characterized transcripts from seven genes that are expressed differently by the worker- and the queen-destined larvae at a critical point in their development. They are indeed genes with phenotypic plasticity.[44]


Altruistic Expression and Social Cues

Let us now consider non-eusocial social organisms. Trivers explained the Vampire bats altruistic behavior based on the ASA assumption.[22] However, we can explain that behavior as well without presuming the existence of distinct selfish/altruistic genotypes.

In vampire bats, hungry individuals often solicit food from those that are better fed. Sometimes, an individual obliges and regurgitates to a soliciting individual—and, sometimes, it steadfastly refuses to share food.

Trivers is right as to the reciprocity of this altruistic behavior. Whether a vampire bat shares blood depends on whether the solicitor has given the donor blood earlier or is likely to give blood to the donor later.[45] The same individual behaves selfish and altruistic depending on social circumstances. This conclusion is compatible with reciprocal altruism.[37] However, as genotypes do not change overnight, it is more plausible for the distinct phenotypes to be due to the plasticity of the same allele.

Male adult olive baboons, Papio anubis, help troupe members depending on whether he has earlier received help from those individuals or they are deemed likely of lending him help in the future.[46]

The behavioral strategy of efficient coercion also supports the plastic phenotypic deployment of a single genotype. In ten studied social insect species, social sanctions kept individuals altruistic in situations where they would have been selfish (Wenseleers and Ratnieks 2006).[47]

Altruists can become selfish. In white-fronted bee-eater, Meropsis bullockoides, young males do not set up nests because older males harass them. They become altruistic helpers instead. However, some of these helpers turn selfish once they build their own nests.[48]

In meerkat societies, the altruistic lower-ranking females become selfish if they attain a higher ranking, not only by breeding but also by killing the infants of other (by then) subordinate females.[49]

Social organisms may assume subordinate roles, not because of their altruistic genes, but because it is the best of the options, they have at the time.[50]

The reviewed examples seem to confirm the hypothesis of altruism being due to a single genotype with plastic phenotype options. An individual will behave altruistically under particular environmental conditions and will respond selfishly when those same cues are absent. As far as this author knows, no evidence has yet proved that only certain individuals can react altruistically or selfishly, given they act in the same scenario.[37]



An Example We All Know—Do Dogs Show Altruistic Behavior?

We know dogs for teaming together against rivals. That is widespread territorial behavior among pet dogs, strays and wild canids. They also defend their household, including humans and other animals. The females bring up and protect their pups at their own expense. These are all examples of behavior we can classify as altruistic. Even considering the difficulties inherent to the concept of altruism, we described, a prudent statement would be that dogs show reciprocal altruistic behavior under the right circumstances, as the ASP model describes.



Conclusion and Perspectives

The altruism paradox is perhaps among the most interesting in ethology (biology) and has, therefore, preoccupied many researchers. It poses highly pertinent questions to enhance our understanding of behavior, genetics and evolutionary theory. Scientists have proposed various models, some of which are more consensually accepted than others. Still, they have all contributed to increasing our global knowledge. The application of the ASP model to behavior promises perspectives far beyond the one we studied in this paper. Perhaps other behaviors that conventional approaches ascribe to distinct phenotypes are but phenotypic plastic expressions of a genotype.



[1] “Altruism.Merriam-Webster. Merriam-Webster, Incorporated. 2017. Retrieved 2017-10-10.
[2] Altruism, Based on the Random House Dictionary (2017). Dictionary.com Unabridged. Random House, Inc. 2017. Retrieved 2017-10-10.
[3] Slote, M.A. 1964. An Empirical Basis for Psychological Egoism, Journal of Philosophy 61: 530-537. Retrieved: 2017-10-10.
[4] Fox, M.W. 1972. Behaviour of Wolves, Dogs, and Related Canids. Harper & Row. ISBN 978-0060113216.
[5] Rogers, L.J.; Kaplan, G.T. 2003. Spirit of the Wild Dog: The World of Wolves, Coyotes, Foxes, Jackals & Dingoes. Allen & Unwin. ISBN 1865086738.
[6] “An amazing True Story—1/24/2008“. The David Sheldrick Wildlife Trust. 2008. Retrieved 2017-10-10.
[7] Darwin, C. 1871. The Descent of Man, and Selection in Relation to Sex (1st ed. ed.). London: John Murray. ISBN 0-8014-2085-7.
[8] Okasha, S. 2009. Biological altruism. In: Zalta EN (ed) The Stanford encyclopedia of philosophy, winter 2009 edn.
[9] Wynne-Edwards, V.C. 1962. Animal Dispersion in Relation to Social Behaviour. Edinburgh: Oliver & Boyd.
[10] Lorenz, K. 1963. Das sogenannte Böse zur Naturgeschichte der Aggression. Verlag Dr. G Borotha-Schoeler (“So-Called Evil, Toward a Natural History of Aggression”).
[11] Hamilton, W.D. 1964. The genetical evolution of social behaviour, I and II. Journal of Theoretical Biology. 7: 1–16, 17–32. doi:10.1016/0022-5193(64)90038-4.
of sociobiology. Q Rev Biol 82:327–348.
[13] Taylor, C., Nowak, M.A. 2007. Transforming the dilemma. Evolution, 61(10):2281–2292.
[14] Allen, B., Traulsen, A., Tarnita, C.E., Nowak, M.A. 2012. How mutation affects evolutionary games on graphs. J Theor Biol 299:97–105.
[15] Nowak, M.A. 2012. Evolving cooperation. J Theor Biol 299:1–8.
[16] Wilson, D.S. 2015. Does Altruism Exist? Culture, Genes, and the Welfare of Others. Yale UP. ISBN: 0300189494.
[17] Maynard Smith, J. 1964. Group selection and kin selectionNature. 201 (4924): 1145–1147. doi:10.1038/2011145a0.
[18] Darwin, C. 1859. The Origin of Species. Reprint: Modern Library, New York 1998.
[19] Fisher, R. 1930. The Genetical Theory of Natural Selection. The Clarendon Press Oxford. OCLC: 18500548.
[21] 1976 July 29, New Scientist, Section: Letters, Haldane, (Letter to the editor from J. Maynard Smith), Quote Page 247, Column 1 and 2, Published by New Science Publications, London; Now Published by Reed Business Information, UK.
[22] Trivers, R. L. 1971. The evolution of reciprocal altruism. Quarterly Review of Biology. 46: 35–57. doi:10.1086/406755.
[23] Rapoport, A. 1966, Two-Person Game Theory: The Essential Ideas. The University of Michigan Press. Ann Arbor, MI,
[24] Mérő, L. 1998. The Prisoner’s Dilemma. In: Moral Calculations. Springer, New York, NY. https://doi.org/10.1007/978-1-4612-1654-4_3.
[25] Fisher, R. 1930. The Genetical Theory of Natural Selection. The Clarendon Press Oxford. OCLC: 18500548.
[26] Zahavi, A. 1975. Mate selection—a selection for a handicap. Journal of Theoretical Biology. 53 (1): 205–214. PMID 1195756. doi:10.1016/0022-5193(75)90111-3.
[27] Zahavi, A. 1977. The cost of honesty (Further remarks on the handicap principle). Journal of Theoretical Biology. 67 (3): 603–605. doi:10.1016/0022-5193(77)90061-3.
[28] Koeslag, J.H. 1997. Sex, the Prisoner’s Dilemma Game, and the evolutionary inevitability of cooperation. Journal of Theoretical Biology. 189: 53–61. PMID 9398503. doi:10.1006/jtbi.1997.0496.
[29] Mech, L.D.; Boitani, L. 2010. Wolves: Behavior, Ecology, and Conservation. Chicago: University of Chicago Press. ISBN 0226516989.
[30] Abrantes, R. 2015. Ethology—The Study of Animal Behavior in the Natural Environment. Cambridge: Wakan Tanka Publishers.
[31] Williams, G.C. 1966. Adaptation and Natural Selection: a Critique of Some Current Evolutionary Thought. Princeton: Princeton University Press.
[32] Dawkins, R. 1976. The Selfish Gene. Oxford: Oxford University Press (1989). pp. 183–185. ISBN 0192860925.
[33] Nowak, M. A.; Tarnita, C. E.; Wilson, E. O. 2010. The evolution of eusociality. Nature. 466 (7310): 1057–1062. doi:10.1038/nature09205. ISSN 1476-4687.
[34] Wilson, E.O. 2012. The Social Conquest of Earth. Liveright. ISBN-10: 978-0871403636/0871403633.
[35] Markvoort, A.J.; Sinai, S., Nowak, M.A.. 2014. Computer simulations of cellular group selection reveal mechanism for sustaining cooperation. Journal of Theoretical Biology 357 (2014) 123–133.
[36] Wilson, E.O. 2005. Kin selection as the key to altruism: its rise and fall. Soc Res 72:159–166.
[37] Yakubu, Y. 2013. The Altruism Paradox: A Consequence of Mistaken Genetic Modeling. Biol Theory (2013) 8:103–113 DOI 10.1007/s13752-013-0120-4.
[38] Pigliucci, M. 2001. Phenotypic Plasticity: Beyond Nature and Nurture. Johns Hopkins University Press, Baltimore.
[39] Kelly, S.A., Panhuis, T.M., Stoehr, A.M. 2012. Phenotypic Plasticity: Molecular Mechanisms and Adaptive Significance. Comprehensive Physiology. pp. 1417–39. doi:10.1002/cphy.c110008. ISBN 978-0-470-65071-4. PMID 23798305.
[40] Winston, M.L. 1987. The Biology of the Honeybee. Harvard University Press, Cambridge, MA.
[41] Prete, F.R. 1990. The conundrum of the honey bees: one impediment to the publication of Darwin’s theory. J Hist Biol 23:271–290.
[42] Albert, S., Bhattacharya, D., Klaudiny, J., Schmitzova, J. & Simuth, J. 1999. The family of major royal jelly proteins and its evolution. J. Mol. Evol. 49, 290– 297.
[43] HGSC The Honeybee Genome Sequencing Consortium. 2006. Insights into social insects from the genome of the honeybee Apis mellifera.
Nature, 443, pages 931–949 (2006)
[44] Evans, J.D., Wheeler, D.E. 1999. Differential gene expression between developing queens and workers in the honey bee, Apis mellifera.
Proc Natl Acad Sci USA 96:5575–5580.
[45] Wilkinson, G.S. 1984. Reciprocal food sharing in the vampire bat. Nature 308:181–184.
[46] Packer, C. 1977. Reciprocal altruism in olive baboons. Nature 265:441–443.
[47] Wenseleers T., Ratnieks, F.L.W. 2006. Enforced altruism in insect societies. Nature 444:50.
[48] Emlen, S.T., Wrege, P.H. 1992. Parent–offspring conflict and the recruitment of helpers among bee-eaters. Nature 356:331–333.
[49] Young, J.A., Clutton-Brock, T. 2006 Infanticide by subordinates influences reproductive sharing in cooperatively breeding meerkats. Biol Lett 2:385–387.
[50] Gadagkar, R. 1997. Survival strategies: cooperation and conflict in animal societies. Harvard University Press, Cambridge, MA.

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Do Non-Human Animals Have Consciousness?

Do Non-Human Animals Have Consciousness?


Do non-human animals have consciousness? Does this question belong to philosophy, ethology, neurobiology or physics?

Many philosophers distinguish between awareness and consciousness, awareness being a form of perception, and consciousness involving a special kind of self-awareness. Consciousness, in this view, requires a propositional awareness that it is I who am feeling or thinking. However, a dissociation between conscious and unconscious perception can occur in people with brain damage, who yet can make correct judgments even though they are not conscious of what they see.

The question of animal consciousness is indeed a difficult one. The spectrum of scientific opinion is vast. Some believe that consciousness does not occur in animals, and others maintain that most animals have consciousness. The difficulty of arriving at an acceptable and consensual definition of consciousness confounds the situation further.

To define consciousness in such a way that it only fits Homo sapiens sapiens seems to me too anthropocentric. To argue that if we have it, they can’t have it, is definitely committing the fallacy of anthropodimorphism, the opposite of anthropomorphism (Abrantes 2017).

Our tentative definition is: “Consciousness is the presence of mental images and their use by the organism to regulate its behavior. To be conscious is to be aware of what one is doing or plans to do. It is having a purpose and intention in one’s actions” (Abrantes 2012). This definition is quasi-identical to McFarland’s (1998).

Yet, we can imagine intentional behavior that does not involve consciousness. The standard examples of automatic, absent-minded activities, are walking and driving, where we solve relatively complex problems apparently without being conscious of (at least) some of the intentional states through which we must be passing.

“Another way to put the point is to note that intentionality alone is not sufficient for introspectibility, and so, insofar as introspectibility in the normal human case is a necessary condition for a state’s being conscious, intentionality is therefore also not sufficient for consciousness” (Rey 2008).

Another definition, proposed by Chandroo et al., is, ” Consciousness might be broadly described as an awareness of internal and external stimuli, having a sense of self and some understanding of ones place in the world” (Chandroo et al. 2004).

Damasio (2010) states that “[…] no one can prove satisfactorily that nonhuman, nonlanguage beings have consciousness, core or otherwise, although it is reasonable to triangulate the substantial evidence we have available and conclude that it is highly likely that they do.”


Private Experiences—Emotions

Emotion has subjective, physiological, and behavioral manifestations that are difficult to reconcile with each other. An emotion is a private experience. There is no way we can know the emotional experiences of another person. We tend to assume they are the same as ours, but we have no experimentally conclusive or logical way of verifying this.

In scientific terms, we cannot assume that animals have particular subjective feelings any more than we are entitled logically to make such assumptions about other people. In physiological terms, emotional states in humans are typically accompanied by autonomic changes, but these are not a reliable guide to identifying particular emotional states. Most animals, at least vertebrates, react to stressors in roughly the same way whether their emotional response is one of fear, of aggression or sexual nature.

Darwin (1872) postulated that facial expressions and other behavioral signs of emotion had evolved from protective responses and other utilitarian aspects of behavior. Darwin’s description was somehow anthropomorphic, which prompted psychologists to react. Lloyd Morgan (1882) advocated an approach devoid of speculation about the private thoughts and feelings of animals. The behaviorist attitude that the private mental experiences of animals cannot be the subject of scientific investigations dominated the first three-quarters of the 20th century.

The behaviorist position seems unassailable, but we can circumvent it in two ways. One argument (the logical one) is that although we cannot prove that animals have subjective experiences, it may be true nevertheless. No proof (yet discovered) does not conclusively imply non-existence. Another (the evolutionary probability argument) is to argue that it is unlikely, from an evolutionary stance, that there should be a marked discontinuity between humans and other animals in this respect. Therefore, if we accept the existence of subjective experiences in humans, then we are forced to admit that animals might have them as well.



Are animals aware of themselves in the sense that they know what posture they are adopting and what action they are taking? Sensory information from the joints and muscles is available to the brain, so it seems that animals should be aware of their behavior.

In an experiment, researchers trained rats to press one of four levers (Beninger et al. 1974). The rats learned to push a different bar, depending on whether they were grooming, walking, rearing up or remaining still when the buzzer sounded. In a sense, the rats must have been aware of their actions, which does not necessarily mean that they are conscious of them. They may be aware of their actions as they are aware of external stimuli.


The MSR (Mirror Self-Recognition) Test

Many animals respond to a mirror as if they saw another member of their species. Does that prove self-awareness? After many years of research, this remains a controversial question. There is evidence that chimpanzees and orangutans can recognize themselves in the mirror. Does the ability to respond to parts of one’s body seen in a mirror indicate self-awareness?

The questions are whether the MSR test is suitable for some species and whether it demonstrates self-awareness. Animals can be self-aware in ways the mirror test cannot measure, e.g., distinguishing between their own and others’ songs or scents (Bekoff 2002). Also, animals can pass the MSR, not necessarily having self-awareness (Cammaerts 2015). Very few species have passed the MSR test (Turner 2015).

The MSR test has limited value when we apply it to species that primarily use senses other than vision, as for example in dogs that mainly use olfaction and audition. Dogs do seem to discriminate their own odor from that of other dogs and to spend more time investigating their own modified odor ‘image,’ precisely as subjects who pass the MSR test do (Horowitz 2017).

A capacity for self-recognition in a mirror does not necessarily imply an awareness of one’s own psychological states and the understanding that others possess such states (Povinelli, 1998).



Does the ability to imitate the actions of others indicate self-awareness? Imitations do not demonstrate the implication of mental states because of the extensive training involved in the experiments.

We take the ability to imitate as a sign of intelligence. Parrots, Psittaciformes, and mynah birds, Sturnidae, can reproduce human sounds with extraordinary fidelity. Are they particularly intelligent?

To be able to imitate, an animal must perceive the external auditory or visual example and match it with a set of motor instructions of its own. For example, a baby who imitates an adult waving must somehow associate the sight of the hand with his own motor instructions for waving. The baby does not need to be aware that he has a hand, it merely has to connect a particular perception with a specific set of motor commands. How that is done, it is a mystery, but the question of whether imitation necessarily involves self-awareness is debatable.

Even though deliberately copying behavior would be a strong argument for self-awareness, we cannot be sure all apparently imitative behavior is.

Allelommetic behavior (synchronous behavior, mimetic behavior, imitative behavior, and social facilitation) may have evolved because a specific synchronization was advantageous. Sometimes, environmental cues initiate this behavioral synchrony (seen in dogs, horses, sheep, chicken, etc.) (Miller 1996, Stoye, S. et al. 2012).

Humans often find themselves assuming similar postures without being aware of that (not conscious of that). It may be the result of empathy, which may also have developed in some species because of the conferred benefits.

Does the brain potential associated with movement occur before or after we are aware of our movement intention? Do I think, “I’m going to move my finger” and then do it? Or does it happen the other way round: I move my finger, and then I’m aware of that? Does consciousness have a causal influence on movement decision? (Guggisberg et al. 2013).


Awareness of Others—Empathy

Empathy means some awareness of others as beings with feelings similar to our own. Some researchers argue that the evolution of close-knit societies made recognition of others, i.e., empathy, advantageous. Empathic behavior is subject to evolutionary laws as any other behavior.

Bischoff-Köhler (1990) investigated the onset of empathy in infants. The results showed that between 16 and 24 months of age, there was a transition from non-recognition to mirror recognition and a simultaneous transition from non-empathy to empathy. Moreover, these transitions occurred at the same age in a given child.

The nature of the phenomenon of empathy in animals is also a topic of investigation (Preston & de Waal 2002). Mice that observe a cagemate in pain are more sensitive to painful stimuli than mice that see an unfamiliar mouse similarly treated (Langford et al. 2006). Empathy can also be the best explanation for some elephant behavior (Byrne et al. 2008).

Finally, empathy does not need to be an all-or-nothing phenomenon (DeWall 1996). Neither does consciousness. As there are various degrees of empathy, there are possibly different levels of consciousness.

The concrete encounter of self and other fundamentally involves empathy, as a unique and irreducible kind of intentionality. Thus, empathy seems to be a precondition for consciousness (Thompson 2001).

However, self-awareness (consciousness) does not always require empathy. That means we can conclude that animals showing empathy must be self-aware. However, we cannot conclude that those who lack manifestations of empathy do not possess self-awareness.


Pain—Do Animals Have to be Conscious in Order to Suffer?

It is difficult to define and analyze pain as the interpretation of findings rest primarily on the behavioral criterion we use. A pure withdrawal reflex would probably not be a good indication of pain. Such reflexes are widespread in the animal kingdom, occur in very primitive animals, and are not always associated with any strong aversive.

The criterion of crying out in pain is not good either. While a dog or a monkey scream in pain when they are seriously injured (or even less seriously), an antelope torn to pieces by a predator remains relatively silent.

Do animals have to be conscious in order to suffer? When we are unconscious, we do not suffer pain or mental anguish because parts of our brain are deactivated. However, we do not know whether these parts are involved only in consciousness or also in other aspects of brain activity. Thus, we cannot say that because we do not experience pain when we are unconscious means that consciousness and suffering are intrinsically related. Perhaps whatever makes us unconscious also stops the pain, but the two are not causally connected.

The truth is that we have no conception of what the conscious experiences of animals might involve if they exist because we have no precise understanding of what consciousness is. Therefore, we can draw no conclusions about the relationship between consciousness and suffering in animals.

Amidst our ignorance, it would be wrong to assume that suffering in animals is confined to those that are intelligent, that use language or that show evidence of conscious experience.

Orch OR Consciousness

Orch-OR is fully compatible with the view that non-human animals possess consciousness to some degree or another. In fact, the opposite would be absurd (illustration from “Consciousness in the Universe: A Review of the ‘Orch OR’ Theory.” Physics of Life Reviews, 2014).

Consciousness as a Quantum State—Orchestrated Objective Reduction (Orch OR)

The Orch-OR model, based on quantum physics, suggests that consciousness originates from microtubules and actions inside neurons (Hameroff 1988, Hameroff and Penrose 2016).

Classic and quantum physics differ in their accounting of events. When I hit a pool ball, I use traditional physics (and geometry) to predict where it will be at any particular moment. I expect it to do so (assuming I hit it correctly). However, in quantum physics, such expectation is null and void. According to the (Copenhagen) interpretation of quantum mechanics, any movement is unknown until it is observed. That is not as weird as it might seem. Imagine, I close my eyes just before I hit the cue ball. While my eyes remain shut, the object ball is both pocketed and non-pocketed. It is first when I open them, that the ball is definitely in one place. Physicists refer to this observation, which determines what happened, as a wave collapsing into a single state.

In quantum systems, inside the neuron, Hameroff and Penrose argue that it is every single collapse of the wave function that returns a conscious moment. Their model met some criticism. Most scientists believe the brain is too warm and wet for quantum states to have any influence on neuronal activity because quantum coherence only seems possible in fairly shielded and cold environments. Biological processes, in general, seem too messy for quantum physics o thrive.

However, researchers have recently found that quantum effects are indeed significant for particular biological processes, like photosynthesis (Engel 2007, Brookes 2017). When a photon hits an electron in a leaf, the electron delivers it to another molecule (the reaction center), which converts light into chemical energy (and feeds the plant). The electron uses the quantum effect of superposition, where a particle can be in two places at once while testing various routes to the reaction center where the photosynthesis occurs. Then, it takes the most efficient one. Besides photosynthesis, olfaction may also be a product of quantum processes (Brookes 2017).

Another support for the idea that quantum physics are indeed possible in the inhospitable organic environment of the brain is via the Phosphorus molecule. The central idea is that the Phosphorous molecule in the brain with its nuclear spin can potentially act as a qubit (quantum bit) and promote quantum computation. This hypothesis circumvents the problem of quantum decoherence by proposing that the qubits remain stable, in spite of the higher temperature of the brain, by organizing themselves into a Phosphate ring (Fisher 2015).

The microtubules, essential in the Orch-OR model, may very well be the first cause of thought. The traditional view is that neurons fire when a channel within the cell membrane opens, flooding the neuron with positively charged ions. Once a determined threshold is reached, an electrical signal travels down the axon—the nerve fibers within the neuron—and the neuron fires. Axons connect neurons to other cells, and inside each axon are nanowires, including the microtubule. Bandyopadhyay found that he could apply a charge to the microtubule, causing activity to raise in the neuron. The nanowires fire thousands of times faster than the average activity in a neuron. The neuron, opposite prevailing scientific knowledge, wasn’t the first cause of the human thought process (Bandyopadhyay 2014).

According to the Penrose–Hameroff model, consciousness results from discrete physical events; such events have always existed in the universe as non-cognitive, proto-conscious events. Biology evolved a mechanism to orchestrate such events and to pair them to neuronal activity, resulting in meaningful, cognitive, conscious moments of quantum state reduction. In the Orch OR theory, these conscious events are terminations of quantum computations in brain microtubules reduced by objective reduction (OR), and having experiential qualities. In this view, consciousness is an intrinsic feature of the action of the universe (Hameroff 1998, Hameroff & Penrose 2016).

Darwin’s theory of natural selection suggests that life evolved by natural selection in incremental steps and random mutations. Therefore, we would not expect the substantial level of coherence across the brain that would be necessary for the non-computable Orch OR of conscious human understanding to appear any other way. More primitive (less elaborated) forms must have preceded it, shown variation, and been subject to natural selection. Thus, proto-conscious Orch OR states might have emerged step by step in the course of evolution.

Orch-OR is fully compatible with the view that non-human animals possess consciousness to some degree or another. In fact, the opposite would be absurd.

Quantum Consciousness Abrantes Billiards

When I hit the cue ball, I use traditional physics (and geometry) to predict where it will be at any particular moment. I expect it to do so (assuming I hit it correctly). However, according to the Copenhagen interpretation of quantum mechanics, any movement is unknown until it is observed. Imagine, I close my eyes just before I hit the cue ball. While my eyes (and my opponent’s, Michael McManus, in this case) remain shut, the object ball is pocketed and non-pocketed at once. It is first when we open our eyes, that the object ball is definitely pocketed.

The Cambridge Declaration on Consciousness

Publicly proclaimed in Cambridge, UK, on July 7, 2012, at the Francis Crick Memorial Conference on Consciousness in Human and non-Human Animals, reads:

“The absence of a neocortex does not appear to preclude an organism from experiencing affective states. Convergent evidence indicates that non-human animals have the neuroanatomical, neurochemical, and neurophysiological substrates of conscious states along with the capacity to exhibit intentional behaviors. Consequently, the weight of evidence indicates that humans are not unique in possessing the neurological substrates that generate consciousness. Non-human animals, including all mammals and birds, and many other creatures, including octopuses, also possess these neurological substrates” (Low, Philip, et al. 2012).

The essential point in this declaration is that concurrent studies concluded that human and some non-human animals share the neurological substrates that we consider indispensable to generate consciousness. That may indeed be helpful in our quest for non-animal consciousness.

Otherwise, the document (as other similar ones) is more a political statement than scientific proof. The fact that it is signed by many (argumentum ad populum) prominent (argumentum ad verecundiam) persons does not add anything to the truth or falsity of its statement about consciousness (Copi 2014). Were the opinion of many the truth, the earth would be flat, after having been created in seven days (Genesis 1 and 2). Were the opinion of prominent persons the truth, DNA would have three intertwined strands (Linus Pauling), and life could originate from inanimate matter (Aristotle); and the Spiroptera carcinoma would (falsely) cause cancer (for which, in 1926, Johannes Fibiger won the Nobel Prize in Medicine).


Final Note

Human consciousness is by definition subjective and private. We access it through verbal, non-verbal, and instrumental records. Animals do not have language (as we define it), but we can still study their consciousness via behavioral investigations, as we do in preverbal infants. Like humans, animals display different behaviors depending on levels of consciousness. During sleep or anesthesia, no individual—unconscious or having low levels of consciousness—independently of species, can process information (not the full range, at least). On the other hand, behavioral and neurobiological data lead us to the conclusion that animals can express some forms of what we call a higher level of consciousness.

The subject of non-human animal consciousness is relevant to many topics, e.g., ethics and theory of mind. Some scientists and philosophers believe that the foundations have been set for addressing (at least) some of the questions about animal consciousness in an empirically way. Some remain skeptical, maintaining that subjective phenomena are beyond the reach of scientific research. The arguments on both sides are many, and the jury is still out.



As a sort of conclusion, it is the view of this author that we have two main unsolved problems related to consciousness: (1) to formulate a conclusive and clear operational definition, and (2) to devise a valid verification method for single species.

As such, at this moment, the most prudent statement seems to be that animals (humans included) show varying degrees of consciousness depending on species. It appears beyond any reasonable doubt that some have it and, therefore, (1) if some have it, others (high) probably have it, too, albeit differently—and (2) that some have it, doesn’t necessarily imply all have it—unless, of course, Hameroff and Penrose are right.

Thank you to John Larsen and Parichart Thongparkdee Abrantes for the exciting exchange of ideas we have had while I wrote this paper.


Abrantes, R. 2012. Ethology—The Study of Animal behavior in the Natural Environment. Wakan Tanka Pub. Online flipping-page book.
Abrantes, R. 2017. Do Animals Have Feelings? Ethology Institute Website. 

Bandyopadhyay, A. 2014. Evidence of massive global synchronization and the consciousness. Physics of Life Reviews.
Bekoff, M. (2002). Animal reflections. Nature. 419 (6904): 255. doi:10.1038/419255a.
Bekoff, M. and Sherman, P.W. 2004.Reflections on animal selves. TRENDS in Ecology and Evolution Vol.19 No.4 April 2004.

Beninger et al. 1974. The ability of rats to discriminate their own behaviours. Canadian Journal of Psychology (Revue Canadienne de Psychologie) 28(1):79-91 DOI: 10.1037/h0081979
Bischof-Köhler, D. (2012). Empathy and Self-Recognition in Phylogenetic and Ontogenetic Perspective. Emotion Review, 4(1), 40–48.
Brown 2015. Brown, C., 2015. Fish intelligence, sentience and ethics. Animal Cognition, 18(1): 1-17.
Brookes, J.C. 2017. Quantum effects in biology: golden rule in enzymes, olfaction, photosynthesis and magnetodetection. Proc Math Phys Eng Sci. 2017 May; 473(2201): 20160822. doi: 10.1098/rspa.2016.0822.
Byrne, R. W., et al. (2008). Do Elephants Show Empathy? Journal of Consciousness Studies, 15(10–11), 204–225.

Cammaerts, M-C, and Cammaerts, R. 2015. Are ants (Hymenoptera, Formicidae) capable of self recognition? (PDF). Journal of Science. 5(7): 521–532.
Chandroo et al. 2004 Chandroo, K.P.; Yue, S.; Moccia, R.D. 2004. An evaluation of current perspectives on consciousness and pain in fishes. Fish and Fisheries, 5(4): 281-295.
Clayton, D.A. 1978. Socially facilitated behavior. Quarterly Review of Biology, 53: 373-392.
Collin, A. and Trestman, M. 2017. Animal Consciousness. The Stanford Encyclopedia of Philosophy (Winter 2017 Edition), Edward N. Zalta (ed.),
Copi, I. 2014. Introduction to Logic. Pearson Education Limited. ISBN 10: 1-292-02482-8.

Damasio, A.R. 2010. Self comes to mind: constructing the conscious brain (PDF). New York: Pantheon Books, 2010. ISBN-10: 150124695X

Darwin, C. 1859. On the origin of species by means of natural selection, or The preservation of favoured races in the struggle for life. London: J. Murray.
Darwin, C. 1871). The descent of man and selection in relation to sex. London; Princeton, NJ: J. Murray ; Princeton University Press, 1981.
Darwin, C. 1872. The expression of the emotions in man and animals. London; Oxford: J. Murray ; Oxford University Press, 1998.
Dawkins, M.S., 1998. Through our eyes only? The search for animal consciousness. Oxford University Press, 1998. ISBN-10: 0198503202
Dawkins, M. S. 2012. Animal Suffering: The Science of Animal Welfare. Springer Science & Business Media. ISBN 9400959052, 9789400959057.
Dennett, D. C. (1969). Content and Consciousness—An Analysis of Mental Phenomena. London: Routledge and Kegan Paul. ISBN: 9780710065124
Dennett, D.C. 1991. Consciousness Explained (PDF). New York: Back Bay Books/Little, Brown and Company, 1991. 511 p.
Dennett, D.C. 1995. Darwin’s Dangerous Idea—Evolution and the Meanings of Life. New York: Simon & Schuster. ISBN: 0-684-80290-2. http://www.inf.fu-berlin.de/lehre/pmo/eng/Dennett-Darwin%27sDangerousIdea.pdf.

Engel, G.S. et al. 2007. Evidence for wavelike energy transfer through quantum coherence in photosynthetic systems. Nature volume 446, pages 782–786 (12 April 2007).

Fisher, M. P.A. 2015. Quantum cognition: The possibility of processing with nuclear spins in the brain. Annals of Physics 362 (2015) 593–602.

Gallego, M.B. 2011. The bohm-penrose-hameroff model for consciousness and free will theoretical foundations and empirical evidences. Pensamiento 67 (254):661-674
Gallup, G. G. Jr. 1970. Chimpanzees: Self-Recognition (PDF). Science, 167 (3914), 86-87.
Gallup, G. G. Jr. 1982. Self‐awareness and the emergence of mind in primates. American Journal of Primatology, 2(3), 237–248.
Guggisberg et al. 2013. Timing and awareness of movement decisions: does consciousness really come too late? Front Hum Neurosci. 2013;7:385.

Hameroff, S.R. 1998. Quantum computation in brain microtubules? The Penrose–Hameroff ‘Orch OR’ model of consciousness. Phil. Trans. R. Soc. Lond. A (1998) 356, 1869–1896
Hameroff, S.R. and Penrose, R. 2016. Consciousness in the Universe: An Updated Review of the “ORCH OR” Theory. In Biophysics of Consciousness: A Foundational Approach by R. R. Poznanski, J. A. Tuszynski and T. E. Feinberg. World Scientific, Singapore. (pp 520-630).
Horowitz, A. 2017. Smelling themselves: Dogs investigate their own odours longer when modified in an “olfactory mirror” test. Behavioural Processes. 143C: 17–24. doi:10.1016/j.beproc.2017.08.001. PMID 28797909.
Hume, D. 1888. A Treatise of Human Nature, edited by L.A. Selby-Bigge (PDF). Oxford: Oxford University Press. ISBN: 9780198245889.
Huxley, T. H. 1874. On the hypothesis that animals are automata, and its history. Fortnightly Review, 95, 555-580.

James, W. 1879. Are we automata? Mind, 4, 1–22. Online.

Langford, D. et al. 2006. Social Modulation of Pain as Evidence for Empathy in Mice. Science 312(5782):1967-70. DOI: 10.1126/science.1128322.
Libet B., Gleason C. A., Wright E. W., Pearl D. K. 1983. Time of conscious intention to act in relation to onset of cerebral activity (readiness-potential). The unconscious initiation of a freely voluntary act. Brain 106, 623–642 10.
Low, Philip et al. 2012. The Cambridge Declaration on Consciousness Publicly proclaimed in Cambridge, UK, on July 7, 2012, at the Francis Crick Memorial Conference on Consciousness in Human and non-Human Animals.

Nagel, T. 1979. Mortal questions. Cambridge. Cambridge University Press (1613). ASIN: B074RB5VTD.

Neindre, P. et al. 2017. Animal consciousness. European Food Safety Authority. 2017, 165 p.

McFarland, D. 1998. Animal Behaviour: Psychobiology, Ethology and Evolution. Benjamin Cummings; 3 ed. ISBN-10: 0582327326.
Morgan, C. L. 1882. Animal intelligence. Nature, 26, 523-524.
Miller, R.M. 1996. Allelomimetic behavior. Journal of Equine Veterinary Science. 16(7): 282–284.

Povinelli, D.J. 1998. Can animals empathize? Maybe not. Scientific American, pp. 67–75.
Preston, S. and de Waal F.B.M. 2002. Empathy: Its ultimate and proximate bases. Behav. and Brain Science (2002) 25, 1–72.

Rey, G. 2008. Even Higher-Order) Intentionality Without Consciousness. Revue internationale de philosophie 2008/1 (n° 243), pages 51 à 78. 

Stoye, S. et al. 2012. Synchronized lying in cattle in relation to time of day. Livestock Science. 149(1–2): 70–73.

Thompson, E. 2001. Empathy and Consciousness(PDF). Journal of Consciousness Studies, 8, No. 5–7, 2001, pp. 1–32
Turner, R. 2015, 10 Animals with Self Awareness. Retrieved 23 November 2015.

de Waal, F. (1996), Good Natured: The Origins of Right and Wrong in Humans and Other Animals. Cambridge, MA: Harvard University Press.

Zahavi, D. (2011). Empathy and direct social perception. Review of Philosophy and Psychology, 3, 541–558.

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“Please” in Animal Language

Please In Animal Language

Saying ‘please’ to your animal can make the whole difference between success and failure. The question is, how do we say that in animal language?

Simplicity is a virtue in life as well as in science and communication. We should always keep that in mind as our animal training gets going.

We can argue for using the dog’s name because it is the simplest signal for us; consequently, we should give the dog a simple and short name. “Adventurous Beautiful Sunset Over the Hills” is undoubtedly a poetic name with its grace if you are inclined to this kind of verse. It looks good in a pedigree but far too complicated for any practical use. It is harder to remember, than a nickname like ‘Bongo,’ it takes longer to pronounce, and it is more difficult to perceive in less than favorable environments. Human nicknames exist for the same reason.

The simplicity of signals is a principle that we should always remember when we plan the training of an animal. All signals should have simple forms, no matter whether they are acoustic, visual or tactile.

Imagine that we are in the same room and consider the following example. I tell you, “please, come here.” The objective of my signal is to have you to move to where I am. ‘Come’ means, “move or travel toward or into a place thought of as near or familiar to the speaker,” or in simpler words, “move to me.”
The addition of ‘here’ is superfluous. ‘Here’ is where I am. If I did not want you to come to where I am, I would not say, ‘come,’ I would say ‘go.’
‘Please’ is in a sense also superfluous. It does not add anything to the behavior you must perform. We use it as a matter of convention because we somehow lost some of our ability to communicate by other than verbal signals. I say, ‘please’ to set you in the right emotional frame of mind to comply with my signal, but I could do that as well without using it. If I said to you, ‘come’ with a smile in my face, a twinkle in my eye and a gentle tone in my voice, I would achieve the same and maybe even better.

“Please” in animal language is not a question of words.

Even though it seems undoubtedly easier, if arguably poorer, to use common words to elicit emotions in our human interactions, it is impossible to accomplish the same when communicating with an animal. There is no way we can explain to an animal what we want to achieve with ‘please.’

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When communicating with an animal, we are better off by just using ‘come’ instead of “please, come here.” It is simpler and conveys to the animal all it needs to know. We can also choose to use ‘here.’ It has the same qualities and none of the mentioned disadvantages. The emotional function of ‘please’ in animal language is better substituted by friendly body language, facial expression, and tone of voice, which are easily detectable by a social animal. ‘Please’ might also influence your state of mind—you are friendlier when you say ‘please’ than when you do not—but here you must compromise with the animal’s innate characteristics. It is easier for the animal to understand a bodily or tonal ‘please’ than a verbal.

Finally, there are situations when we do not need to use ‘please’ and others where we achieve better our goal without it; the same goes for our communication with animals. Sometimes, we will need to use a more assertive body language, facial expression, and tonal voice to achieve our objective; and yet other times we need to be very assertive.

We must assess any particular situation and decide how to modulate our signals. There are two elements in a signal: (1) the factual, which is an operant controlled by the consequences and (2) the emotional, which is the respondent and which the signal itself elicits. It is our job to control both so that we achieve the desired goal, and there is no magical formula to do so.

The factual part of it is clear. We only have to know the science behind it and comply with its rules. It is the part you can learn in the course “Animal Training My Way–Merging Ethology and Behaviorism.” The emotional part, which deals with empathy, is a difficult one. Either you have it, or you don’t. You may acquire it through experience, or you may not, and no one can help you with that.

This article is an excerpt from Roger Abrantes’ book, “Animal Training My Way—Merging Ethology and Behaviorism,” included in the course “Ethology and Behaviorism.

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Evolutionarily Stable Strategies and Behavior

Evolutionarily Stable Strategies and Behavior (DovesAndHawks).

Evolutionary biologists imagine a time before a particular trait existed. Then, they postulate that a rare gene arises in an individual and ask what circumstances would favor the spread of that gene throughout the population. If natural selection favors the gene, then the individuals with the genotypes incorporating that particular gene will have increased fitness. A gene must compete with other genes in the gene pool and resist any invasion from mutants,to become established in a population’s gene pool.

In considering evolutionary strategies that influence behavior, we visualize a situation in which changes in genotype lead to changes in behavior. By ‘the gene for sibling care,’ we mean that genetic differences exist in the population such that some individuals aid their siblings while others do not. Similarly, by ‘dove strategy,’ we mean that animals exist in the population that do not engage in fights and that they pass this trait from one generation to the next.

At first sight, it might seem that the most successful evolutionary strategy will invariably spread throughout the population and, eventually, will supplant all others. While this does occur, it is far from always being so. Sometimes, there is no single dominant strategy. Competing strategies may be interdependent in that the success of one depends upon the existence of the other and the frequency with which the population adopts the other. For example, the strategy of mimicry has no value if the warning strategy of the model is not efficient.

Game theory belongs to mathematics and economics, and it studies situations where players choose different actions in an attempt to maximize their returns. It is a good model for evolutionary biologists to approach situations in which various decision makers interact. The payoffs in biological simulations correspond to fitness—comparable to money in economics. Simulations focus on achieving a balance that would be maintained by evolutionary strategies. The Evolutionarily Stable Strategy (ESS), introduced by John Maynard Smith in 1973 (and published in 1982), is the most well known of these strategies. Maynard Smith used the hawk-dove simulation to analyze fighting and territorial behavior. Together with Harper in 2003, he employed an ESS to explain the emergence of animal communication.

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An evolutionarily stable strategy (ESS) is a strategy that no other feasible alternative can better, given that sufficient members of the population adopt it. The best strategy for an individual depends upon the strategy or strategies which other members of the same population adopt. Since the same applies to all individuals in that particular population, a mutant gene cannot invade an  ESS successfully.

The traditional way to illustrate this problem is the simulation of the encounter between two strategies, the hawk and the dove. When a hawk meets a hawk, it wins on half of the occasions, and it loses and suffers an injury on the other half. Hawks always beat doves. Doves always retreat against hawks. Whenever a dove meets another dove, there is always a display, and it wins on half of the occasions. Under these rules, populations of only hawks or doves are no ESS because a hawk can invade a population made up entirely of doves and a dove can invade a population of hawks only. Both would have an advantage and would spread in the population. A hawk in a population of doves would win all contests, and a dove in a population of hawks would never get injured because it wouldn’t fight.

However, it is possible for a mixture of hawks and doves to provide a stable situation when their numbers reach a certain proportion of the total population. For example, with payoffs as winner +50, injury -100, loser 0, display -10, a population consisting of hawks and doves (or individuals adopting hawk and dove strategies) is an ESS whenever 58,3% of the population are hawks and 41,7% doves; or when all individuals behave at random as hawks in 58,3 % of the encounters and doves in 41,7%. The percentages, or point of equilibrium, depend on costs and benefits (or the pay-off, which is equal to benefits minus costs).

Evolutionarily stable strategies are not artificial constructs. They exist in nature. The Oryx, Oryx gazella, have sharp pointed horns, which they never use in contests with rivals and only in defense against predators. They play the dove strategy. Up to 10% per year of Musk Ox, Ovibos moschatus, adult males die as a result of injuries sustained while fighting over females. They play the hawk strategy.

An ESS is a modified form of a Nash equilibrium. In most simple games, the ESSes and Nash equilibria coincide perfectly, but some games may have Nash equilibria that are not ESSes. Furthermore, even if a game has pure strategy Nash equilibria, it might be that none of those pure strategies are ESSes. We can prove both Nash equilibria and ESS mathematically (see references).

Peer-to-peer file sharing is a good example of an ESS in our modern society. BitTorrent peers use Tit for Tat strategy to optimize their download speed. Cooperation is achieved when upload bandwidth is exchanged for download bandwidth.

Evolutionary biology and sociobiology attempt to explain animal behavior and social structures (humans included), mainly in terms of evolutionarily stable strategies.


Featured image: The traditional way to illustrate Evolutionarily Stable Strategies is the simulation of the encounter between two strategies, the hawk and the dove.

Learn more in our course Ethology. Ethology studies the behavior of animals in their natural environment. It is fundamental knowledge for the dedicated student of animal behavior as well as for any competent animal trainer. Roger Abrantes wrote the textbook included in the online course as a beautiful flip page book. Learn ethology from a leading ethologist.


Are You Teaching Your Pet Superstitious Behavior?


Superstitious behavior is behavior we erroneously associate with particular results. Animals create superstitions as we do. If by accident, a particular stimulus and consequence occur a number of times temporarily close to one another, we tend to believe that the former caused the latter. Both reinforcing and inhibiting consequences may create superstitious behavior. In the first case, we do something because we believe it will increase the odds of achieving the desired result (we do it for good luck). In the second case, we do not do something because we do not want something else to happen (it gives bad luck).

In 1948, B.F. Skinner recorded the superstitious behavior of pigeons making turns in their cages and swinging their heads in a pendulum motion. The pigeons displayed these behaviors attempting to get the food dispensers to release food. They believed their actions were connected with the release of food, which was not true because the dispensers were automatically programmed to dispense food at set intervals.


Some cases of CHAP (Canine Home Alone Problems) could be superstitious behavior. The dog believes that if it barks long enough at the door, someone will open it because it has happened before. Many CHAP cases are not even remotely connected with anxiety as the dog owners erroneously presume.

Superstitious behavior is extremely resistant to extinction. Skinner found out that some pigeons would display the same behavior up to 10,000 times without reinforcement. Displaying a behavior expecting a reinforcer, and receiving none, increases persistence. It’s like we (as well as other animals) feel that if we continue long enough the reinforcement will follow sooner or later.

As always, being an evolutionary biologist, the first question that comes to my mind is, “what conditions would favor the propagation of superstitious behavior?” Making correct associations between events confers a substantial advantage in the struggle for survival. That is what understanding (or adapting to) one’s environment means. The benefits of getting one association right outweigh the costs of making several wrong associations, so much that natural selection favors those who tend to make associations rather than those who do not—and that’s why superstitious behavior is highly resilient to extinction.

Featured image: Warning: superstitious behavior is easy to create and extremely difficult to extinguish.

The Function of Champing Behavior

The Function of Champing Behavior

Champing (or chomping) is a noisy chewing motion, despite there being nothing to chew. This behavior is associated with friendliness, pacifying of an opponent, insecurity, or submission, depending on degree and context.

There is a pacifying element in all forms of champing. Pacifying behavior (Latin pacificare, from pax = peace and facere, facio = to make) is all behavior with the function of decreasing or suppressing an opponent’s aggressive or dominant behavior or restoring a state of tranquility. Licking, nose poking, muzzle nudging, pawing, yawning, and twisting are common pacifying behaviors that dogs offer one another and us.

Champing is a behavior widely used by canines in situations ranging from mild unease to more severe concern.

Champing is one of the first sounds that puppies hear—their sibling’s suckling. It is, therefore, a sound associated with satisfaction. Redirection of the champing behavior assumes later a pacifying function—attempting to turn an unpleasant situation into a pleasant one. Initially, the pups connect camping with the appeasement of hunger.


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).

Champing is a straightforward and efficient way to show friendliness towards a dog. Curiously, this behavior appears to have a relaxing effect on most mammals. Newborn mammals suckle and connect sucking sounds (chomping) with pleasant and desirable consequences. Jane Goodall points out that she used to break a twig and pretend to champ it to pacify chimpanzees showing some unease. I often use chomping when in the presence of dogs and horses showing some degree of distress.

Featured image: Champing behavior has a pacifying function—attempting to turn an unpleasant situation into a pleasant one.

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Learn more in our course Ethology. Ethology studies the behavior of animals in their natural environment. This is a fundamental course for the serious student of animal behavior as well as animal trainers. Roger Abrantes wrote the textbook included in the online course as a beautiful flip page book. Learn ethology from a leading ethologist.


Signal and Cue—What is the Difference?

MaleLionAndCubChitwaSouthAfricaLucaGaluzzi2004 (Signal and Cue—What is the Difference?)

In the behavioral sciences, there is some confusion about the meaning of the terms signal and cue (as with so many other terms) and some authors use it interchangeably. To make it even more difficult, communication theory also uses the same terms with slightly different meanings and in the theatre and movies world a ‘cue’ is actually a ‘signal.’

However, in behavioral sciences, the general consensus (see references below) is that signal and cue have the following meanings.

signal is a perceivable behavior or feature that has evolved and has acquired the specific characteristic of conveying information about the signaler or the signaler’s environment. Information (communication) changes the behavior or the beliefs of the receiver.

This definition of signal implies that if a signal changes the behavior of an organism, this change of behavior must be profitable to both sender and receiver more often than not, or otherwise, signalers would cease to send the signal and receivers would cease to respond. This definition distinguishes, in principle, a signal from coercion, although some signals may be coercive, e.g. threats.

In general, signals must be honest and reliable, or otherwise they cease to have any effect (receivers don’t behave appropriately) and they undermine communication (honest senders will not benefit from sending the signals). However, some signals can tolerate a certain degree of dishonesty, all depending on the costs and benefits for all parties. H. W. Bates discovered in 1861 that some (palatable) butterflies had an advantage in mimicking (Batesian mimicry) poisonous butterflies, which is detrimental to the poisonous butterflies inasmuch as it turns their signals of unpalatability less reliable. On the other side, some species use the same signals to convey the same information and they all benefit from it (Mullerian mimicry).

cue is any feature that an organism can use as a guide to display a particular behavior or series of behaviors. The classical example is the mosquito seeking a mammal to bite and flying up wind when it detects CO2. The CO2 is a cue for the mosquito, but it is surely not a signal sent by the mammal, which would prefer to remain undetected and not be bitten. Intentionality is the key element to distinguish signals from cues.

A cue is a regularity, a pattern that either is permanently ‘on,’ or is ‘on’ and ‘off” depending on specific conditions, e.g. a rock, a tree, or the position of the sun in the sky cues us of directions, and dark clouds cues us of impending rain. The rock, the tree, the sun and the clouds are not there to give us information, but they do if we interpret them correctly. A signal is more malleable, more intentional and we can turn it ‘on’ and ‘off’ in response to relevant cues in the environment, e.g. the warning cry that many species (signal) issue in response to the appearance (cue) of a feared predator.

Cues are traits or actions that benefit the receiver exclusively. The sun and the rock do not profit from us getting our bearings. When a mouse by accident makes a rustling sound in the leaves and attracts a predator increasing the risk of being killed, the sound is a cue for the predator about the location of its prey. When an alert animal deliberately gives a warning call to a stalking predator resulting in the predator giving up the hunt, this sound, the alert call, is a signal both for conspecifics and the predator. Different species can, thus, communicate by means of signals which both recognize and behave accordingly.

Secondary sexual traits are features that distinguish the two sexes of a species, but that are not directly part of the reproductive system. They are probably the product of sexual selection for traits, which give an individual an advantage over its rivals in courtship and competitive interactions. Secondary sexual traits are also cues for the opposite sex. They are not directly related to a better production of offspring, but are normally good indicators of better sperm quality or egg production, e,g, manes of male lions (Panthera leo) and long feathers of male peacocks (Pavo cristatus). In humans, visible secondary sexual traits include enlarged breasts of females and facial hair on males.

The study of signals and cues is more complex that it may appear at first sight. Cues can become signals. In 1952, Niko Tinbergen described ritualization as the evolutionary process whereby a cue may be converted into a signal, e.g. the canine behavior of baring teeth. In 1975, Zahavi described the handicap principle where the reliability of some signals is ensured because they advertise greater costs than absolutely necessary, e.g. the exaggerated plumage of the peacock.

We must understand correctly what the intentionality of signals means and not to confound the intentionality of the signal itself with its origin, development and evolution. Signals do not origin by design with a determined purpose. Some features or behaviors just happen at a certain time to be efficient for an organism in generating in another organisms the right behavior at the right time. If they convey an advantage to these organisms in their struggle for survival (and reproduction), they will spread in the population (provided these organisms reproduce). With time, they gain intentionality and become true signals, but their origin was accidental like everything else. This is the reason why I had to modify (some extensively) the definitions I use in this text and I had to create new ones—to make them compatible with the Darwinian theory of evolution.

Applying the principle of simplicity, as always, I suggest the following definitions:

signal is everything that intentionally changes or maintains the behavior of the receiver. A cue is everything that unintentionally changes or maintains the behavior of the receiver (see “The 20 Principles All Animal Trainers Must Know“).

These definitions open for the possibility to better distinguish between the intentional signals (proper signals) we send and the unintentional ones (which are cues). For example, many dog owners say “no” to their dogs meaning “stop what you are doing,” but their (unintentional) body language (cue) says “yes.”

In conclusion: signal is the most correct term to denominate what we use when we communicate with our animals; and signals may assume many forms, auditory (the words we use), visual (the hand movements and body language we use), olfactory (in canine detection work), tactile (a touch, very common in horse training) and probably also palatable.

So, enjoy the consequence of your (intentional) signals and be careful with any cues you may be (inadvertently) sending to your favorite animal. Enjoy as well your further studies of this fascinating topic: animal communication.


References and further readings

  • Dawkins, M. S., and T. Guilford (1991). The corruption of honest signalling. Animal Behaviour 41:865–873.
  • Donath, J. (2007). Signals, cues and meaning (February draft for Signals, Truth and Design. MIT Press)
  • Hasson, Oren (1997). Towards a general theory of biological signaling. Journal of Theoretical Biology 185: 139-156.
  • Hauser, Marc D. and Mark Konishi, eds. (1999). The design of animal communication. Cambridge: Bradford/MIT Press.
  • Maynard Smith, John and David Harper (1995). Animal signals: Models and terminology. Journal of Theoretical Biology 177: 305-311.
  • Maynard Smith, John and David Harper (2003). Animal signals. Oxford University Press, UK.
  • McFarland, D. (1999). Animal Behaviour. Pearson Education Limited, UK.
  • Otte, D. (1974). Effects and functions in the evolution of signaling systems. Annual Review of Ecology and Systemat- ics 5:385–417.
  • Saleh, N et al. (2007) Distinguishing signals and cues: bumblebees use general footprints to generate adaptive behaviour at flowers and nest. Arthropod-Plant Interactions, 2007, 1:119–127
  • Schaefer, H. M. and  Braun, J. (2009). Reliable cues and signals of fruit quality are contingent on the habitat in black elder (Sambucus nigra). Ecology, 90(6), 2009, pp. 1564–1573.
  • Searcy, W. A., and S. Nowicki (2005). The evolution of animal communication. Princeton University Press, Princeton, New Jersey, USA.
  • Tinbergen, N. (1952). The curious behavior of the stickleback. Scientific American December 1952.
  • Zahavi, A. (1975). Mate selection: a selection for a handicap. Journal of Theoretical Biology 53:204–214.

Featured image: Secondary sexual traits, as the mane of the male lion, are powerful cues (Photo by Luca Galuzzi via Wikipedia).

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Animal Learning

Should We Reinforce the Effort or the Result?


The main difficulty in some learning processes is to reinforce the right behavior at the right time, which bad teachers, bad parents, and bad trainers do not master (bad means inefficient, and it is not a moral judgment).

If you ask, “should we reinforce the effort or the result?” you are liable to get as many answers supporting the one opinion as for the other. Supporters of the effort system sustain that reinforcing results creates emotional problems when one doesn’t succeed and decreases the rate of even trying. Advocates of the result method defend that reinforcing the effort encourages sloppiness and cheating.

I shall argue in the following for and against both theories and prove that it is not a question of either/or, rather of defining clearly our criteria, processes, and goals.

I shall compare the learning of some skills in dogs and humans because the principles are the same. The difference between them and us is one “of degree, not of kind,” as Darwin put it.

I will use SMAF to describe some processes accurately where I find it advantageously. If you are not proficient in SMAF, and you’d like to be, please read “Mission SMAF— Bringing Scientific Precision Into Animal Training.”

Much of my personal work with dogs (and rats and Guinea Pigs) is and has been detection work, mainly narcotics and explosives, but also person search, tobacco, and other scent detection work. One of the first signals I teach the animals is a disguised reinforcer.

With dogs, I use the sound ‘Yes’ (the English word). The signal part of this signal/reinforcer means, “continue what you’re doing,” and the reinforcer part, “we’re OK, mate, doing well, keep up.” That is a signal that becomes a reinforcer: Continue,sound(yes) that becomes a “!+sound”(yes).

The difference between the most used “!±sound”(good-job) and “!+sound”(yes) is that the former is associated and maintained with “!-treat”(small food treat) and “!-body(friendly body language); and the latter with a behavior that will eventually produce “!-treat”. The searching behavior does not provide a treat, but continuing searching will eventually (find or no find). That is why “!+sound”(yes) is a disguised Continue,sound(yes) or the other way around.


Search’ means “Go and find out whether there is a thing out there.” The signal ‘Search’ (Search,sound) does not mean ‘Find the thing.’ Sometimes (most of the time) there’s nothing to find.

Why do I need this interbreeding between a signal and a reinforcer?
Because the signal ‘Search’ (Search,sound) does not mean ‘Find the thing.’ Sometimes (most of the time) there’s nothing to find, which is good for all of us (airports and the likes are not that full of drugs and explosives).

So, what does Search,sound mean? What am I reinforcing? The effort?
No, I’m not. We have to be careful because if we focus on reinforcing the effort, we may end up reinforcing the animal just strolling around, or any other accidental or coincidental behavior.

I am still reinforcing the result. ‘Search’ means “Go and find out whether there is a thing out there.” ‘Thing’ is everything that I have taught the dog to search and locate for me, e.g., cocaine, hash, TNT, C4.

“Go and find out whether there is a thing out there” leaves us with two options equally successful: ‘here’ and ‘clear.’ When there is a thing, the dog answers ‘here’ by pointing at its apparent location (I have taught it that behavior). When there is no thing, that is precisely what I want the animal to tell me: the dog answers ‘clear’ by coming back to me (again because I have taught it that). We have two signals and two behaviors:

Thing,scent => dog points (‘here’ behavior).
∅Thing,scent => dog comes back to me (‘clear’ behavior).

The signals are part of the environment. I do not give them, which does not matter: a signal (SD) is a signal.(1) An SD is a stimulus associated with a particular behavior and a particular consequence or class of consequences. When we have two of them, we expect two different behaviors, and when there is none, we expect no behavior. What fools us, here, is that, in detection work, we always have one and only one SD, either one or the other. Having none is impossible. Either we have a scent, or we don’t, which means that either we have Thing,scent or we have ∅Thing,scent, requiring two different behaviors as usually. The one SD is the absence of the other.

Traditionally, we don’t reinforce a search that doesn’t produce a positive indication. To avoid extinguishing the behavior, we use ‘controlled positive samples’ (a drug or an explosive, we know it is there because we have placed it there to give the animal a possibility to obtain a reinforcer).

That is a correct solution, except that it teaches the dog that the criterion for success is ‘to find’ and not ‘not to find,’ which is not true. ‘Not to find’ (because there is nothing) is as good as ‘to find.’ The tricky part is, therefore, to reinforce the ‘clear’ and how to do it to avoid sloppiness (strolling around) and cheating.

Let us analyze the problem systematically.

The following process does not give us any problems:

Search,sound => Dog searches => “!+sound”(yes) or Continue,sound(yes) => Dog searches => Dog finds thing (Thing,scent) => Dog points (‘here’ behavior) => “!±sound”(good-job) + “!-treat”.

No problem, but what, then, when there is no thing (∅Thing,scent)? If I don’t reinforce the searching behavior, I might extinguish it. In that situation, I reinforce the searching with “!+sound”(yes):

“Search,sound” => Dog searches => “!+sound”(yes) => Dog searches => ∅Thing,scent => Dog comes back to me (‘clear’ behavior) => “!±sound”(good-job). */And I can also give “!-treat”*/

Looks good, but it poses us some compelling questions:
How do I know the dog is searching versus strolling around (sloppiness)?
How do I know I am reinforcing the searching behavior?

If I reinforce the dog coming back to me, then, next time I risk that the dog will take a quick round and get to me right away: that is the problem. I want the dog to return to me only when it finds nothing (the same as didn’t find anything).

To reinforce the searching behavior.
To identify the searching behavior versus strolling around (sloppiness). How can I make sure that the dog always searches and never only rambles around?

Reinforcing the searching behavior with “!+sound”(yes) works. OK.

Remaining problem:
I have to reinforce the ‘clear’ behavior (coming back to me), but how can I make sure that the dog always searches and never strolls around (avoid sloppiness)?
How can I make sure that the dog has no interest in being sloppy or cheating me?

To teach the dog that reinforcers are available if and only if:
1. The dog finds the thing. Thing,scent => Dog sits => “!±sound”(good-job) + “!-treat”.
2. The dog does not ever miss a thing. ∅Thing,scent => Dog comes back to me => “!±sound”(good-job) + “!-treat”.

I teach the dog gradually to find things until I reach a predetermined low concentration of the target scent (my DLO—Desired Learning Objective). In this phase of training, there is always one thing to find. After ten consecutive successful finds (my criterium and quality control measure), all producing reinforcers for both the searching (“!+sound”(yes)) and the finding (“!+sound” + “!-treat”), I set up a situation with no thing (∅Thing,scent). The dog searches and doesn’t find anything. I reinforce the searching and the finding (no-thing) as previously. Next set-up, I make sure there is a thing to find, and I reinforce both searching and finding.

I never reinforce not-finding a thing that is there or finding a thing that is not there (yes, the last one is an apparent paradox).

Consequence: the only undesirable situations for a dog are: (1) not-finding a thing that is there (the dog did not indicate Thing,scent), or (2) indicating a thing that is not there (the dog indicates ∅Thing,scent).

(1) Thing,scent => Dog comes back to me (‘clear’ behavior) => [?±sound] + [?-treat].
(2) ∅Thing,scent => Dog points (‘here’ behavior) => [?±sound] + [?-treat].

That is (negatively) inhibiting negligence, but since it proves to increase the intensity of the searching, we cannot qualify it as an inhibitor. Therefore, we call it a non-reinforcer: “∅±sound”, “∅-treat”.
In the first case:

Thing,scent => Dog comes back to me => [?±sound] + [?-treat].
Thing,scent => Dog comes back to me => “∅±sound”, “∅-treat”.
Thing,scent => Dog comes back to me => “∅±sound”, “∅-treat” => Dog searches (more intensively) => Thing,scent => Dog points (‘here’ behavior) => “!±sound” + “!-treat”.

In the second case, I have to be 100% sure that there is indeed no-thing. The training area must be free of any scent remotely similar to the scent we are training (Thing,scent). Particularly in the first phases of the training process, this is imperative, and a trainer who misses that is committing major negligence.

Should the dog, nevertheless, show ‘here’ for ∅Thing,scent, then we can use the same procedure as above:

∅Thing,scent => Dog shows ‘here’ behavior => “∅±sound”, “∅-treat” => Dog searches (more intensively) => ∅Thing,scent => Dog comes back to me (‘clear’ behavior) => “!±sound” + “!-treat”.

What if later the dog doesn’t find a thing that is there in a lower concentration than the one I used for training, or masked by other scents?

No problem—that is not the dog’s fault. I didn’t train it for it. The dog doesn’t know that it is committing a mistake by giving me a (wrong) ‘clear.’ As far as the dog is concerned, the room is clear. For the dog, it is a ‘clear’: ∅Thing,scent => Dog comes back to me => “!±sound” + “!-treat”. The dog was not strolling around and is not cheating me.

Comparing to humans:
I reinforce the behavior of the child trying to solve a math problem. Yes, we must always reinforce (or inhibit) a behavior, not the individual. “Well done, but you got it wrong because…” The solution may be incorrect, but the method was correct. Then, it is all a question of training. More or better training will eliminate the ‘wrong.’ Maybe, it was caused by a too abrupt increase in the difficulty curve of the problem (which is the teacher’s problem). We are not reinforcing trying; we are reinforcing the correct use of a method (a desired process).

Why reinforce the process?
We must reinforce the process because of its emotional consequences. The dog and the child must accept the challenge, must want to be tried and to be able to give their best in solving a problem.

Are we reinforcing the effort rather than the success?
No, we are not. Reinforcing the effort rather than the result can and will lead to false positives. The animal indicates something that it is not there because it associates the reinforcer with the behavior, not the thing. Children give us three-four consecutive, quick and wrong answers if we reinforce the trying, not the process (thinking before answering).
We reinforce the result (success) only. When the dog doesn’t find because there’s nothing to find, that is a success. When the dog doesn’t find because the concentration was too low, that is a success because ‘too low’ is here equal to ‘no-thing.’ When the child gets it wrong, it is because the exercise exceeded the actual capacity of the child (not trained to that). No place to hide for trainers, coaches, teachers, and parents.

We are still reinforcing success and exactly what we trained the dog and the child to do. We don’t say to the child, “Well, you tried hard enough, good.” We say, ” Well done; you did everything correctly. You just didn’t get it right because you didn’t know that x=2y-z and you couldn’t know it.” Next time, the child gets it right because now she knows it; and if not, it is because x=2y-z exceeds the capacity of that particular child, at that particular moment, in which case, there’s nothing to do about it.

The same with the dog: the dog (probably) will not indicate 0.01g of cocaine because I trained it to go as low as 0.1g. When I reinforce the dog’s ‘clear,’ I say, “Well done, you did everything correctly, you just didn’t get it right because you didn’t know that 0.01g cocaine is still the thing.” Now, I train the dog that ‘thing’ means ‘down to 0.01g cocaine’ and either the dog can do it or it cannot. If it can, good. If it cannot, there’s nothing we can do about it.

We reinforce result, success, not the effort, not trying. We must identify success, have clear criteria for success, plan a progressive approach to our goal, a gradual increase in difficulties. We must be able to recognize limits and limitations in ourselves, in the animal species with which we work, in the individuals we tutor, in the particular skill we teach. We must know when we cannot improve a skill any further and when someone, human or not, cannot give us more than what we get; and be satisfied with that.
1 Strictly speaking, the scent, which the detection dog searches, is not a signal, but a cue, because it is not intentional. In this context, however, it is an SD because we have conditioned it to be so, and we can, therefore, call it a signal. Please, see “Signal and Cue—What is the Difference?

Featured image: Learning is a complex process The main difficulty in some learning processes is to reinforce the right behavior at the right time, which bad teachers, bad parents, and bad trainers do not master. We must reinforce the process because of its emotional consequences. The dog and the child must accept the challenge, want to be challenged, to be able to give their best in solving the problem, not giving up.

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The 20 Principles The 20 Principles All Animal Trainers Must Know explains clearly and concisely how we best can train animals independently of species—dog, horse, cat, Guinea pig, marine mammal, etc. This course is a 'must' for all animal trainers.

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Learn more in our course Canine Scent Detection, which will enable you to pursue further goals, such as becoming a substance detection team or a SAR unit. You complete the course by passing the double-blind test locating a hidden scent. You take the theory online in the first three lessons. In lesson four, you train yourself and your dog, step by step until reaching your goal. We will assign you a qualified tutor to guide you, one-on-one, either on-site or by video conferencing.

Canine Scent Detection

Laughter is the shortest distance…


As you have figured out by now, I enjoy finding proof that humans are not that different from other forms of life. We share many characteristics with the other living creatures on our blue planet. Today, I have one more example for you—laughter.

Laughing is an involuntary reaction in humans consisting of rhythmical contractions of the diaphragm and other parts of the respiratory system. External stimuli, like being tickled, mostly elicit it. We associate it primarily with joy, happiness, and relief, but fear, nervousness, and embarrassment may also cause it. Laughter depends on early learning and cultural factors.

The study of humor and laughter is called gelotology (from the Greek gelos, γέλιο, meaning laughter).

Chimpanzees, gorillas, bonobos, and orangutans display laughter-like behavior when wrestling, playing or tickling. Their laughter consists of alternating inhalations and exhalations that sound to us like breathing and panting.

Rats display long, high frequency, ultrasonic vocalizations during play and when tickled. We can only hear these chirping sounds with proper equipment. They are also ticklish, as are we. Particular areas of their body are more sensitive than others. There is an association between laughter and pleasant feelings. Social bonding occurs with the human tickler, and the rats can even become conditioned to seek the tickling.*

A dog’s laughter sounds similar to a regular pant. A sonograph analysis of this panting behavior shows that the variation of the bursts of frequencies is comparable with the laughing sound. When we play this recorded dog-laughter to dogs in a shelter, it can contribute to promoting play, social behavior, and decrease stress levels.*

Victor Borge once said, “Laughter is the shortest distance between two people.” Maybe, it is simply the shortest distance between any two living creatures.


* Panksepp & Burgdorf, 2003, ‘Laughing’ rats and the evolutionary antecedents of human joy?; Simonet, Versteeg & Storie, 2005, Dog-laughter: Recorded playback reduces stress related behavior in shelter dogs.

Featured image: We laugh, but we are not the only ones.

Bonding with Your Dog—Are You Doing it Properly?

Bonding with Your Dog (Desi and Dog)

Bonding with Your Dog—Are You Doing it Properly? Bonding in animal behavior is a biological process in which individuals of the same or different species develop a connection. The function of bonding is to facilitate co-operation.

Parents and offspring develop strong bonds so that the former take care of the latter and the latter accept the teachings of the former. This serves both parties best. As a result of filial bonding, offspring and parents or foster parents develop an attachment. This attachment ceases to be important once the juvenile reaches adulthood, but may have long-term effects upon subsequent social behavior. Among domestic dogs, for example, there is a sensitive period from the third to the tenth week of age, during which normal contacts develop. If a puppy grows up in isolation beyond about fourteen weeks of age, it will not develop normal relationships.

Males and females of social species develop strong bonds during courtship motivating them to care for their progeny, so they increase their chances of the survival of 50% of their genes.

Social animals develop bonds by living together and having to fend for their survival day after day. Grooming, playing, mutual feeding, all have a relevant role in bonding. Intense experiences do too. Between adults, surviving moments of danger together seems to be strongly bonding.

Bonding behavior like grooming and feeding seems to release neurotransmitters (e.g., oxytocin), which lowers innate defensiveness, increasing the chances of bonding.

We often mention bonding together with imprinting. Even though imprinting is bonding, not all bonding is imprinting. Imprinting describes any type of phase-sensitive learning (learning occurring at a particular age or a specific life stage) that is rapid and (apparently) independent of the consequences of behavior. Some animals appear to be preprogrammed to learn about certain aspects of the environment during particular sensitive phases of their development. The learning is pre-programmed in the sense that it will occur without any apparent reinforcement or punishment.

Our dogs in our domestic environments develop bonds in various ways. Grooming, resting with each other, barking together, playing and chasing intruders are strong bonding behaviors. Their bonding behavior is by no means restricted to individuals of their own species. They bond with the family cat as well and with us, humans.

Bonding is a natural process that will inevitably happen when individuals share responsibilities. Looking into one another’s eyes is only bonding for a while, but surviving together may be bonding for life—and this applies to all social animals, dogs and humans included.

We develop stronger bonds with our dogs by doing things together rather than by just sitting and petting them. These days, we are so afraid of anything remotely connected with stress that we forget the strongest bonds ever originate under times of intense experiences. A little stress doesn’t harm anyone, quite the contrary. I see it every time I train canine scent detection. The easier it is, the quickest it will be forgotten. A tough nut to crack, on the other hand, is an everlasting memory binding the parties to one another.

I even suspect one of the reasons we have so many divorces these days is that we want everything to be easy, and oh so pleasant, that in the end, nothing is holding the two together—but that’s another story for maybe another time.

Featured image: Bonding develops stronger and more readily in stressful situations. SAR handlers and their dogs have probably some of the strongest bonds we witness between the two species. Photo: Désirée Mallè, Alpine Rescue Team, and her dog.

Learn more in our course Ethology and Behaviorism. Based on Roger Abrantes’ book “Animal Training My Way—The Merging of Ethology and Behaviorism,” this online course explains and teaches you how to create a stable and balanced relationship with any animal. It analyses the way we interact with our animals, combines the best of ethology and behaviorism and comes up with an innovative, yet simple and efficient approach to animal training. A state-of-the-art online course in four lessons including videos, a beautiful flip-pages book, and quizzes.