Dissecting Behavior

Dogs, Science, and the Biology of Behavior

Bad Science: Quadrants of Operant Conditioning

People become dog trainers for various reasons. Often, these individuals will talk about a dog’s “performance,” yet this undoubtedly has a variety of interpretations. After all, what is performance? Is it speed? Strength? Accuracy? Reliability? Chat up a few trainers involved in any professional sport (canine or human) and you will see that there are numerous beliefs both for which methods produce the best results for the desired performance as well as for what reasons. Should our toes be pointing straight ahead or at an angle when doing a squat? Should we stretch before or after an activity? With dogs though, the question is even more convoluted because here the concerns are not just about performance: they are also about welfare.

Animal welfare is a vast topic and one that cannot be approached from A-Z in a single sitting. Many philosophers and scientists devote their entire lives to traversing the quagmires of non-human animal welfare issues and so I am not going to put all of my roulette chips down on 28 black and defend my choice in the never ending spin of the animal welfare debate wheel. In a perfect world, conversations are always productive. In the actual world, this is often a rare occurrence. But to the point, the conversation that might be the least productive in dog welfare is the assertion that techniques which use positive reinforcement and negative punishment are ethical; while techniques which use positive punishment and negative reinforcement are unethical. This issue is so emotionally charged and emblazoned in the industry that often the supporting evidence for a claim about the ethical nature of a technique revolves solely around the interpretation of what quadrant of operant conditioning the technique relies on.

Skinners-box1-277x300Common illustration of basic learning concepts in a 2×2 grid often called “the Quadrants.”
(note, while often attributed to B.F. Skinner, this is a false attribution)

For example, many trainers claim that a technique called Behavior Adjustment Training (BAT) is unethical because they see it as negative reinforcement. For those unfamiliar, BAT removes a stimulus (which a dog finds threatening) at a distance great enough for the dog to remain calm and not show signs of being overly agonistic (such as growling, snarling, barking, etc.) [note: typically it is the dog that is moving, not the scary stimulus].  Because you repeatedly remove something (in this case, the thing the dog doesn’t “like”) to reinforce calmer behavior, many trainers label this type of training as negative reinforcement—and because negative reinforcement is claimed to be unethical, BAT must therefore be unethical.  Problematically, BAT was designed to steer owners away from using harsh punishments and the method itself creates no signs of undue harm on the dog; so, if the interpretation of the quadrants of operant conditioning cause trainers to conclude that BAT is unethical, then there is a serious problem with the convention because calling BAT unethical is like calling Mr. Snuffleupagus from Sesame Street a serial rapist.

Since 1975, various scientists have pointed out that learning events can rarely, if ever, be labeled solely as positive or negative (e.g. Michael 1975, Baron & Galizio, 2005; Baron & Galizio, 2006; Tonneau, 2007). For example, imagine a rat in a black box at freezing temperature. They have a lever which activates a heater for a short period of time. As the rat stays in the box, an increase (reinforcement) in lever-pressing is noticed over time. Here is the paradox: does lever-pressing increase because of the addition (positive) of heat or the cessation (negative) of cold? The answer is yes.

[This example is paraphrased from an actual experiment conducted by Weiss & Laties published in Science in 1961]

In the physical universe, the addition of one stimulus is always met with the removal of another stimulus. Regardless of what type of matter (energy) this stimulus is, energy cannot be created or destroyed, and so within any closed system you have to remove something to add something and you have to add something to remove something. This is a fundamental property of the universe and is analogous to the idea that two opposing baseball teams cannot win the same game: in order for one team to win, another team has to simultaneously lose.  This prompts us to ask two questions: 1) are the quadrants of operant conditioning mutually exclusive?; and 2) if they are not mutually exclusive, then are we able to stipulate that they are not occurring at the same time during a learning event?

Most examples of what dog trainers consider positive reinforcement rely significantly on negative reinforcement elements (e.g. the removal of hunger). Food is great, but as a motivator we are removing hunger (negative reinforcement), however it is also positive reinforcement for the obvious reason that we are adding food.  This might seem unimportant for the lives of most dogs who are fed to the point of obesity, however in behavior research, most animals are deprived of food before reinforcement begins in learning paradigms, therefore the contingency of food as positive reinforcement is being given to an animal deprived of enough food prior to testing to cause a 15% decrease in body mass.  For perspective, imagine a 180-pound male losing 27 pounds before being handed a cheeseburger as reinforcement and you might appreciate how removing deprivation is not only perhaps a better description of the actual science of reinforcement but also a significant motivator for a rat to start pressing levers in their black box.

A classic example used popularly in psychology textbooks is the example of an aspirin as a negative reinforcement. The idea is that the removal of a headache might increase future aspirin-taking behavior, thus the removal of the aversive headache could be said to increase the frequency of the behavior—or more concisely, the aspirin is negatively reinforcing aspirin-taking behavior. However, we are adding aspirin to the system, so what do we say about the addition of a stimulus that causes the removal of another stimulus that overall causes a consequence which increases or decreases the frequency of the antecedent behavior? Vis-à-vis “aspirin-logic,” the addition of food that removes the feeling of hunger would have to be negative reinforcement as well.  By now it should be clear that there is no mutual exclusivity to the reasoning behind the popular interpretations of the quadrants of operant conditioning, and therefore any conclusion that relies on such a demarcation is neither logical nor scientific.  Simply put, analyzing behavior with a system that relies on the Tweedledee-Tweedledum characterization of reinforcement and punishment (Marr, 2006) in a universe that is beholden to the conservation of energy is a product of improper, massive oversimplification.

It should be appreciated that the difficulty of negotiating positive versus negative effects within a system is common to science.  For example, biologists that enjoy old-fashioned terminology will describe the movement of an organism in relationship to a stimulus a “taxis;” positive taxis is therefore movement toward a stimulus, while negative taxis is movement away from a stimulus. If the reference point for the behavior is the change to the environment (e.g. the appearance of a prey animal) then naturally we would instinctively describe the motion of a predator towards the prey as positive taxis. However, let us instead change the animal to an herbivore like an elk. Imagine a large group of elk munching away on some delicious savory grass. Overtime, the elk wear down the presence of grass in the area they are feeding and they then move toward another area which has more grass present. Are the elk moving toward an area of more food to forage on or away from an area of less food to forage on (i.e. is it positive taxis toward new grass or negative taxis away from no grass)?

It is important to remember that much of what we use to categorize nature are simply conventions, and sometimes their creation is no more sophisticated than what one person decided while reading the latest issue of Science while sitting on the can. One of my favorite illustrations of the sometimes arbitrary nature of conventions is in the way physicists describe torque motions. In physics, a torque that generates movement counterclockwise is notated with a positive force and a torque that generates movement clockwise is notated with a negative force. Why? Because if you replicate the motion of an object moving counterclockwise with your fingers on your right hand, your thumb is pointing up, and if you replicate the motion of an object moving clockwise with your fingers on your right hand, your thumb is pointing down. For this reason it is called the thumb rule.

ImageDespite the overwhelming issue, research papers and essays are still frequently published describing events that are “positive reinforcement” or “negative reinforcement,” therefore this is by no means just a dog industry issue.  Furthermore, responses to these criticisms fail in addressing the issue head on, are unable to provide sound counterarguments, and/or fall back on the pragmatic argument: “well, we don’t have anything better so it is better than nothing.” There are a couple problems with the pragmatic argument. First, define what is “better?” Quadrants create a paradigm view that cannot be supported without the existence of quadrants, so if “better” requires a convention that maintains the theory-laden beliefs of operant conditioning then I would say the pragmatists are correct, just like creationism cannot exist without a God who created the universe as the central hypothesis.  If “better” requires only the need to describe learning events then the pragmatists are definitively wrong because the concepts of reinforcement and punishment are descriptive enough in and of themselves as positive/negative distinctions always have to be clarified further with methodological explanation.

But all of this side steps the heart of the issue: harsh punishment creates the negative and deleterious results we are familiar with because of the threat it presents to the organism.  The ethics here are measured through actual harm, not through the way an animal learned something. Indeed, many dogs might not learn much of anything that is objectively quantifiable in an operant classification after being swung around on a choke chain in a helicopter swing or kicked in the ribs, thus we couldn’t say these events belong to any quadrant because we have to first establish the learned behavior that is operating on the environment.

Ethics does not have a quadrant. It is a complex web of issues that are rarely cut and dry and conversations about dog training through positive and negative quadrant distinctions only obfuscate the discussion at hand. Kicking a dog is unethical because it is harmful and cruel: not because it is “positive punishment.” Dangling a dog in the air as it suffocates is unethical because it too is harmful, cruel and abusive. You cannot design an experiment to show that the Yankees won is true but the Red Sox lost is false in the same way it is impossible to falsify whether it is the addition of a treat or the removal of hunger acting during a learning event. Pragmatists will say “oh, whatever, it’s not a big deal because I know the difference.” Problematically, it’s not only unhelpful to the conversation but it is also unscientific. Science is falsifiable, if it is not, it is not science.


Baron, A., & Galizio, M. (2005). Positive and negative reinforcement: Should the distinction be preserved? The Behavior Analyst / MABA, 28(2), 85–98.

Baron, A., & Galizio, M. (2006). The distinction between positive and negative reinforcement: Use with care. The Behavior Analyst, 29(1), 141.

Marr, M. J. (2006). Through the Looking Glass: Symmetry in Behavioral Principles? The Behavior Analyst, 29(1), 125.

Michael, J. (1975). Positive and Negative Reinforcement, a Distinction That Is No Longer Necessary; Or a Better Way to Talk about Bad Things. Behaviorism, 3(1), 33–44.

Tonneau, F. (2007). Behaviorism and Chisholm’s Challenge. Behavior and Philosophy, 35, 139–148.

Weiss, B., & Laties, V. G. (1961). Behavioral Thermoregulation. Science, 133(3464), 1588–1588. doi:10.1126/science.133.3464.1588

What is behavior?

The Question of Behavior

What exactly is behavior?  What is it that we are examining when we inquire about the behavior of animals?  This question is not as straightforward as one might think.  There is a significant amount of disagreement on the subject that benefits from consideration.  In a paper by Levitis et al. (2009), the question of ‘what is behavior?’ was asked of professors and professionals from three different scientific societies.  Of the eight operational definitions the authors proposed from various texts, the highest agreement on any single definition only included 44% of the respondents.  Even more problematic is that 52% of the respondents showed contradictory answers—implying that the respondents’ conceptions of behavior was not guided by an operational definition of behavior but of their personal biases (most likely due to their individual areas of interest).  For instance, even though 99.1% of the respondents agreed that “geese flying in a V formation” constitutes behavior, 31% of the respondents answered that only individuals, not groups, are capable of behavior.  In argument form, this line of deduction would look like this:

Premise 1:       Only individuals, not groups, are capable of behaving
Premise 2:       Geese flying in a V formation constitute a group
Therefore:       The group of geese is behaving

This is a flawed argument to say the least. The benefit of operational definitions is that they allow us to organize our observations through logical lines of reasoning, thus contradictions in logic are a significant red flag that bias has taken place of a scientific definition.  One potential explanation for this is that in Levitis et al.’s (2009) background research to gather published definitions for the term “behavior,” over one hundred publications that they felt should have had a definition for behavior had completely avoided defining the term, either by choice or due to the assumption that people intrinsically know what behavior is.  I hope it has been successfully illustrated that the latter conclusion is clearly not the case.

The Definition of Behavior

A quick thumb in the dictionary informs us about a common preconception: “behavior is the way in which an animal or person acts in response to a particular situation or stimulus.”  In fair agreement with this definition, Tinbergen defined behavior as “the total movements made by the intact animal” (Tinbergen, 1955).  However, what if we were to talk about the behavior of a plant?  Some species of orchids (e.g. Ophrys speculum) use chemical signals to mimic virgin females of their insect pollinators to deceive male insects into attempting reproduction with their flowers: thus causing the male insect to pollinate the orchid (Ayasse et al., 2003). Deception (the action of altering the perception of another to believe something that is not true in order to gain some personal advantage) is most certainly a phenomenon most anyone would associate with behavior.  However a plant is not an animal, so evidently the concept of behavior has to expand beyond just animals if it is going to include deception.


Ophrys speculum evolved features which mimic female reproductive organs in order to deceive male insects, thus increasing their ability to be pollinated.

Conceivably then we could expand the parameter of the definition to multi-cellular organisms.  Having said that, such a classification would then exclude bacteria.  Bacteria are single-cell organisms that meet all eight of the requirements that define life: (1) they have an internal organization; (2) they store and retrieve information through the organization of nucleotides; (3) they have a metabolism that converts energy from an unusable to a usable form; (4) they respond to the environment; (5) they grow and develop; (6) they regulate and maintain homeostasis; (7) they reproduce; and finally, (8) they evolve.  One could argue that if you momentarily exclude reproduction and evolution, all of the requirements of life facilitate just one organismic phenomenon: homeostasis.  So the first argument from the skeptic must be to show that a living entity that is responding to its environment is not capable of behavior.

It might seem unorthodox for many readers to discuss the behavior of bacteria, however even Skinner was not without his philosophical views of the behavior of single-cell organisms and discussed the stretching of an amoeba’s limbs as a an example of early behavioral evolution (Skinner, 1984).  In fact, amoebas are part of the group Protozoa which comes from the Greek for “first animal.”  Protozoa were named this because of their animal-like behavior, from their movement to their reliance on consuming other organisms for survival.  While I share no disagreement with Skinner regarding the behavior of single cell organisms, where I do disagree is in the view that the work of Jacques Loeb (1915) is sufficient in providing a sound argument that the behavior of “lower” organisms only needs explication in what the organism does as a whole—a recurrent theme which Skinner also applied to the behavior of larger organisms as well (Skinner, 1987).

This view is incredibly limiting.  In the hundreds years following Loeb, we have discovered more about the behavior of all organisms (prokaryotes [archaea & bacteria] and eukaryotes) by investigating internal cellular functions.  Indeed, researchers recently discovered that bacteria are able to activate a host body’s immune response to attack an invading virus that threatens the survival of their host they are living symbiotically with (Ichinohe et al., 2011).  Researchers discovered this behavioral response by analyzing signals that lead to the synthesis of mRNA essential to the immune response—a discovery that has launched an entirely new field of inquiry to other ways medicine might fight deadly diseases.  However, this extraordinary level of symbiotic behavior is still poorly understood, so for our purposes let us take a look at a simple environmental response in a single cell of Escherichia coli as an example of adaptive behavior in bacteria.

When E. coli are in a high solute, hypertonic environment, their life is instantly endangered by diffusion’s closest relative: osmosis.  Osmosis is the movement of water from an area of higher free energy to an area of lower free energy (Saupe, 2013).  Solutes (dissolved molecules) affect this process by affecting the free energy of the water the same way that rush hour traffic decreases the kinetic energy of a freeway—the more traffic in the solution (presence of solutes), the less available free energy.  If the solutes from the E. coli’s environment diffuse across its membrane through its pores, osmosis might destroy and kill the cell because freely permeable water will pass across the membrane until the free energy inside the cell is equal to that outside.  If the presence of solutes is dense enough, the E. coli will explode and die.

E. coli would not have survived for the eons they have been around for if a handful of solutes made them explode, thus their behavioral response is an extremely effective defensive mechanism rooted in their DNA.  When E. coli senses the presence of solutes, they transmit a signal to begin transcribing and translating sections of DNA into proteins that will block the pores and prevent solutes from diffusing across the membrane—thus maintaining homeostasis.  There is no question that these actions of E. coli are essential for their survival and fit perfectly well along side the adaptive behaviors of plants and animals.

Permitting then that we include single-cell organisms in our definition of behavior, what then do we do about viruses?  Viruses have no cells.  They are purely microscopic packets of genetic code wrapped in a protein shell.  Consequently, viruses lack the essential organelles required for replication, thus in order for a virus to reproduce it literally breaks open the membrane of a host cell by using spikes on its protein shell that act like a medieval battering ram to break down the gates and invade.  This allows the virus to use the mechanisms within the fully functioning host cell to replicate its viral genetic code like nefarious molecular zombies (see Grove & Marsh, 2011).  Interestingly, scientists analyze viruses in terms of their (1) physical structure, (2) proximate causation, (3) temporal effect on the fitness of a host which impacts the virus’s capacity for survival, and (4) evolutionary development—Tinbergen’s four questions of behavior (Erkoreka, 2010; also see Tinbergen, 1963).  If viruses are incapable of behavior, are Tinbergen’s four questions irrelevant?  Such a claim would create an upheaval in the behavioral literature.


Here is another way we might ask ourselves about the behavior of viruses.  Let us for the sake of argument claim that viruses are not capable of behavior.  If this is true and we are to exclude these molecular zombies from the definition of behavior, why then do we commonly say that viruses attack cells?  Are we just attributing the behavioral characteristics of animals to non-animal entities or is it really valid to say that these molecular zombies are truly attacking and threatening the homeostasis of an organism?  Even the dictionary cites a definition for “attack” as the aggressive action of a disease.  Viruses obviously meet this criterion so then are we to exclude the behavior of attacking from the definition of behavior just so we can deprive these mysterious molecules of behavior?  While colloquialisms are generally poor grounds for scientific exploration, in this case I would argue that they bear important consideration to the question at hand.  After all, it would be ignorant to deny that common expressions reflect our experiential perceptions of the world (known in science as empirical observation).

The fact that these molecules are nothing but packets of genetic material and literally straddle the definitional requirements of life yet have actions attributable to behavior quickly leads us into questions about non-living entities—for example, do rocks behave?  If an individual was to pick up a rock and throw it, is the rock behaving?  A definition that relied purely on the reactions of matter to its environment would require us to include every rock on Earth as a behaving entity since rocks are quite literally orbiting the Sun as the Earth moves through our solar system.  Using Newtonian physics, we would stipulate that the rock is exerting a force on the Earth equal to that of the force of the Earth on the rock.  Instead of thinking about a small rock, what if we were to look at hydrogen dioxide (water).  The behavior of water makes it one of the most unique molecules in existence.  Three of waters most important behavioral characteristics (i.e. properties) are that it is a powerful solvent, it sticks to itself because it is polar and thus can generate incredible surface tensions, and it is denser as a liquid than as a solid.  If any one of these three properties were different, life would not exist as we know it today.  Molecules have observable, quantifiable properties which respond to other molecules in the environment, therefore why can’t molecules behave?


One might argue for a definition that stipulated some kind of internal propulsion or energy source which might allow us to exclude a rock.  But would this criterion be valid?  Even though the energy used by animals is converted from an unusable to a useable form through internal metabolic processes, we have already established that the source of all energy for every organism comes from the Sun.  The survival of any animal would be immediately threatened without the energy mitochondria (prokaryotic organisms) produce or the symbiotic benefits offered by gut flora (prokaryotic organisms like E. coli), thus the behavior of animals cannot be observed without the impact of the organisms acting on them.  In a similar fashion, no organism lives in a void.  We shouldn’t forget, as silly as the concept of a rock behaving may seem, all animal behavior is subject to the physical laws of nature.  A bird cannot fly without overcoming the necessity to generate lift, neither could energy be converted to facilitate homeostasis or motion without adhering to the intrinsic properties of atomic theory.  If an owl is diving on a field mouse, prey capture could not occur without the effects of gravity—thus the very action pattern depends on the same phenomenon that determines the travel path of a rock that is thrown.  It should be mentioned that the skill of a bird negotiating with the physical properties of lift is equal to any human pilot, even though the mechanisms are obviously different, and while the Border Collie is unlikely to recite Newton’s laws of motion, they have a skill in predicting the motion of a Frisbee equal to any Ultimate Frisbee competitor.

Any general definition of behavior must allow for all of the nuances mentioned so far if it is to permit our inferences to take into account the immense diversity of behavior in the world.  Thus, returning to Tinbergen, I will modify and broaden his insight and offer that behavior is the total measurement of movements (both internal and external—regardless of physical or biological limitations) of an entity through its environment.  More simply: behavior is the response of a system to a stimulus.  Therefore, since every particle in the universe is in motion and endlessly responding to stimuli, by this definition we can conclude that everything—even a subatomic particle—is in a constant state of behavior.

Such an incredibly broad definition may at first seem either ridiculous or of little value, yet I would contend that there is nothing more valuable than to question our preconceptions with a logical and unbiased deductive thought process.  There is no reason to exclude rocks as behaving entities in the world, and a rejection of this has more to do with the novelty of it (“well I just don’t like it..”-type of response) than of the reasoning to the question.  Furthermore, I would argue that the benefit of such a definition provides a common concept to be discussed whether the system involves entire populations or just a single atom.

Previously I cited the opinion from Levitis et al’s (2009) study that 31% of participants believed that only individuals, not groups, are capable of behavior.  Problematically, the behavior of the human body is the result of the sum of movement of molecules from within—therefore how can we say that the movement of trillions of molecules inside a single organism is “behavior” yet preclude the sum of a group of geese performing an action (such as flying in a V formation) as a behaving entity?  I have no doubt that a five-minute discussion with a police officer that has had to respond to a riot can attest to the behavioral differences of an angry mob versus that of an angry individual.  The tendency for human nature to warp their perceptions to fit their preconceptions makes the ability to integrate new information very difficult; especially if that information negates beliefs we have held for a long time.  As Tinbergen states:

“… if we overdo this in itself justifiable tendency of making description subject to our analytical aims, we may fall into the trap some branches of Psychology have fallen into, and fail to describe any behaviour that seems ‘trivial’ to us; we might forget that naïve, unsophisticated, or intuitively guided observation may open our eyes to new problems.  Contempt for simple observation is a lethal trait in any science, and certainly in a science as young as ours.”  (Tinbergen, 1963)

A definition for behavior that includes all matter in the universe, while useful for expanding our perceptive lens, may not initially seem to provide a productive answer for our question of what is animal behavior.  If narrow perspectives risk losing the forest through the tree, this might be like losing the forest in the galaxy.  What should be emphasized is that the question of what is behavior is only productive if the nature of the entity exhibiting the behavior is definitively described.  The behavior of organisms is thus our aim, and this behavior is intrinsically tied to the fact that organisms—regardless of their complexity—are bound to their biological nature.


Ayasse, M., Schiestl, F. P., Paulus, H. F., Ibarra, F., & Francke, W. (2003). Pollinator attraction in a sexually deceptive orchid by means of unconventional chemicals. Proceedings of the Royal Society B: Biological Sciences, 270(1514), 517–522. doi:10.1098/rspb.2002.2271

Erkoreka, A. (2010). The Spanish influenza pandemic in occidental Europe (1918-1920) and victim age. Influenza and Other Respiratory Viruses, 4(2), 81–89. doi:10.1111/j.1750-2659.2009.00125.x

Freeman, S., Quiliin, K., & Allison, L. (2013). Pearson – Biological Science (5th edition.). San Fransisco, CA: Benjamin Cummings.

Grove, J., & Marsh, M. (2011). Host-pathogen interactions: The cell biology of receptor-mediated virus entry. The Journal of Cell Biology, 195(7), 1071–1082. doi:10.1083/jcb.201108131

Ichinohe, T., Pang, I. K., Kumamoto, Y., Peaper, D. R., Ho, J. H., Murray, T. S., & Iwasaki, A. (2011). Microbiota regulates immune defense against respiratory tract influenza A virus infection. Proceedings of the National Academy of Sciences, 108(13), 5354–5359.

Levitis, D. A., Lidicker, W. Z., & Freund, G. (2009). Behavioural biologists do not agree on what constitutes behaviour. Animal Behaviour, 78(1), 103–110. doi:10.1016/j.anbehav.2009.03.018

Loeb, J. (1915). On the role of electrolytes in the diffusion of acid into the egg of Fundulus. Journal of Biological Chemistry, 23(1), 139–144.

Saupe, Stephen G. (2013). Letter to the Editor. American Biology Teacher, 75(1), 4-5.  DOI: 10.1525/abt.2013.75.1.2

Skinner, B. F. (1984). The evolution of behavior. Journal of the Experimental Analysis of Behavior, 41(2), 217–221.

Skinner, B. F. (1987). Whatever happened to psychology as the science of behavior? American Psychologist, 42(8), 780–786. doi:10.1037/0003-066X.42.8.780

Tinbergen, N. (1955). The Study of Instinct. Oxford: Clarendon.

Tinbergen, N. (1963). On aims and methods of Ethology. Zeitschrift für Tierpsychologie, 20(4), 410–433. doi:10.1111/j.1439-0310.1963.tb01161.x

Are dogs and wolves the same species?

The question of whether dogs and wolves are members of the same or different species is a controversial one.  To begin with, species classification is a convention used to help aid in our ability to organize nature and it is anything but definitively objective.  This should not decrease the importance of classifying species, but before we begin to try and understand the question, we will benefit from understanding that the nature of the question is very philosophical.  Always keep in the back of your mind that the personal preference of an individual will always be influential in subjective conclusions.  Therefore, to try and be objective about the conversation I would like to discuss the big picture, and in biology, the big picture is always evolution.

Evolution is often described as cumulative processes so slow that they take between thousands and millions of decades to complete (e.g. Dawkins, 1986).  This is only part of the picture.  We certainly have an in-depth archeological fossil record that shows gradual changes in species over millennia (such as the development of feathers in dinosaurs or the eye-migration of flatfish), however biological changes can also happen in the wink of an eye—at least compared to traditionally conceptualized evolutionary timescales.  Most simply, evolution can be defined as change over time.  But what kind of change?  Does any change constitute evolution? Does any duration sufficiently qualify for “time?”  These are important considerations because whatever definition is chosen will create a first premise assumption from which any arguments will flow from—like the way the lens of a camera manipulates light before entering the camera body and forming an image, so too can a first premise assumption influence our perceptions so that our observations fit a desirable theory instead of the natural phenomenon.

Some evolution happens very slowly (such as the previously mentioned examples of feathers in dinosaurs and the eye-migration of flatfish); however, these changes arose most probably due to mutation and sexual selection, not because these changes condoned a functional advantage in evading hazards or finding food.  Most examples of evolution are due to a change in the characteristics of a group that enable it to survive, thus evolution can be viewed in this light as a response to changes in the environment.  Typically, environments change very slowly and significant changes often ride on the back of natural disasters.  The evolution of dinosaurs into birds was due to a two-fold catastrophe.   Approximately 200 million years ago, atmospheric oxygen declined nearly 20% causing one of the largest extinction periods in Earth’s history (Berner et al., 2006).  This killed off an unprecedented amount of land dwelling animals and threatened aquatic living organisms as well.  For example, some species of fish such as tuna evolved ram-air induction (whereby swimming at high velocity forced water across the gills at a higher speed to ensure maximum oxygen diffusion from water).  As if global suffocation wasn’t bad enough, to add insult to injury, an asteroid the size of Manhattan slammed into Mexico just a few millennia later.  These two factors meant that the only dinosaurs which survived were small and could fly—what we call birds.  Predominantly, it is important to remember that changes to the environment are what drive these kinds of selection processes, especially when these changes create significant mortality rates—a concept I will return to later.

The controversy over the classification of dogs and wolves can be seen on numerous levels, but one that stands out for me is the way in which many wolf-dog hybrid enthusiasts are very passionate that the correct term is not “hybrid” but “wolf dog”—since both the dog (Canis lupus familiaris) and the wolf (Canis lupus lupus) are according to some scientists taxonomically sub-species of Canis lupus.  While this is a relatively recent distinction (originally, Carl von Line classified the dog as Canis familiaris, a different species than the wolf) the taxonomic nomenclature does not determine whether the mating of two animals qualifies as a hybrid.  Hybridization is the interbreeding of individuals from genetically distinct populations, regardless of their taxonomic status (Stronen & Paquet, 2013).  Wolves and dogs may be amazingly similar in their genetics, however they are clearly genetically distinct populations (e.g. vanHoldt et al., 2011).


Principal component analysis of all wolf-like canids for the 48K SNP data set: PC1 represents a wild versus domestic canid axis, whereas PC2 separates wolves (n=198) and dogs (n=912) from coyotes (n=57) and red wolves (n=10). Result shows dogs and gray wolves are genetically distinct (Fst=0.165). PC2 in this analysis of the data demonstrate a geographically based population hierarchy within gray wolves and coyotes (vanHoldt et al., 2011)


Plots show ancestry blocks and their assignments for representative individuals of canid populations with average size of blocks, percent ancestry, and number of generations since most recent admixture (t) indicated. Two-ancestor (coyote, A; gray wolf, B) analyses are presented for a Great Lakes wolf from Minnesota (C), a captive red wolf (D), and an Algonquin wolf (E). Three-ancestor analyses (coyote, A; gray wolf, B; dog, F) are presented for a Northeastern coyote from Vermont (G), a Southern coyote from Louisiana (H), and a Midwestern coyote from Ohio (I) – (vanHoldt et al., 2011)

The supposedly infallible “fact” that dogs are descended from wolves took the world by fire with research into mitochondrial DNA and a publication which appeared in Science titled “Multiple and Ancient Origins of the Domestic Dog” (Vila et al., 1997).  In this paper, the authors concluded that dogs were 135,000 years old—a conclusion which is sheer nonsense (Larson, 2011; Larson et al., 2012).  Over the last decade, geneticists have published paper after paper pointing at different dates and different locations for domestication with very little consensus but most supporting the conclusion that dogs are direct descendants of the wolf.  One important reason for this is because the methodology behind examining mitochondrial DNA (mtDNA) has a very debilitating first premise assumption: that the rate of mtDNA mutation is constant in dogs and wolves despite a massive wolf population bottleneck and an exploding dog population.  This is a problem because both of these population effects cause genetic drift.  Imagine if you take a population and reduce it to a mere handful.  How do you tell whether you are looking at the first members of a new species or the surviving members after a population endangerment?  Likewise, imagine taking two dogs and deciding you will start your own breed.  If your new breed goes through a population explosion, then their DNA will make up a unrepresentative sample of the historical population (this is called the “founder effect”).

Genetic research is awesome, don’t get me wrong, and it cannot be underappreciated that innovations in genetics have opened up wildly exciting new scientific avenues of investigation into organisms.  However, genetic analysis is relatively new to the question of speciation in the animal kingdom and some insight to the Canis lupus dilemma can be gained by looking at the overall ecology of dogs and wolves instead of just their sequence of nucleic acids.  Research that examines genotypes, high-density single nucleotide polymorphisms, epigenetic methylations, mitochondrial DNA, etc., is literally a whole new world, but it is not the whole picture.  The expression of a plant or animal’s DNA is what creates its phenotype (from morphology to behavior), and it is the phenotype that is thus selected for in the environment and we can learn lots by simply examining the phenotype in and of itself.

As previously mentioned, when two genetically distinct species reproduce the offspring is called a hybrid.  However in animals, hybridization is a pretty big deal.  When sperm meets egg, a zygote is formed, thus ecologists look at both prezygotic (before reproduction) and postzygotic (after reproduction) barriers that make hybridization difficult.  Examples of prezygotic barriers include: habitat isolation (where two species are geographically isolated, sometimes living in the same area but rarely meet), behavioral isolation (where two species do not recognize the signals/mating cues of each other or employ different foraging strategies), temporal isolation (where one species might breed in the spring while another breeds in the fall), mechanical isolation (where the “wedding tackle” of one species doesn’t fit in the “hoo-ha” of another species), and genetic isolation (where the sperm and egg of two species are unable to form a zygote).  Postzygotic barriers include reduced hybrid viability (where hybrids fail to develop or reach sexual maturity), reduced hybrid fertility (e.g. mules are hybrids of horses and donkeys and are all sterile), and hybrid breakdown (where the offspring of hybrids have further reduced viability and/or fertility).

The behavioral isolation of dogs and wolves is astronomical because behaviorally there are almost no commonalities between them.  In fact, leaving dogs aside for a moment, very important behavioral distinctions exist just between different groups of wolves that affect their offspring viability (postzygotic barriers).  For example, one of the most important criteria for mate preference in wolves comes down to hunting strategies: wolves with similar hunting and foraging strategies are more likely to mate and teach these strategies to their offspring.  Foraging behavior is a phenotypical characteristic that plays a major role in determining the ecological niche of a species—so much so that wolves who employ different foraging strategies also display different types of social relationships.

Very few dogs hunt for food.  Even in societies which still use dogs for hunting (such as the indigenous Mayagna people of Nicaragua), dogs rarely make the kill.  Their role in the hunt is to bring an animal to ground and make a loud ruckus until the humans can find it and make the killing blow with their machete.  In this capacity, dogs are pound for pound as efficient as a rifle in bringing in meat for the indigenous people of Nicaragua, and the dogs benefit by being given leftovers (Koster, 2008).  It is certainly true that some dogs (some) opportunistically take down and on occasion eat small animals such as rats, possums, cats, etc.  However, dogs like other scavengers fill an important role in the grand ecological picture regarding the flow of biomass (Wikenros et al., 2013).

Hunting in the wild is simply not an available strategy for dogs to survive.  One important reason for this is that the energy dogs would expend to take down and eat small prey animals would be much greater than the energy gained by hunting them.  This is illustrated with African wild dogs (Lycaon pictus), who pound for pound hunt, kill, and consume more meat than any other predator in Africa—this is not because they are greedy, this is because of their metabolic needs.  When you look at African wild dogs, small prey like African hares make up only an average profitability of 0.6kg per hunt (4.8kg per kilometer chased), whereas Wildebeest weighing approximately 100kg make a profitability of 35.2kg per hunt (51kg per kilometer chased).  African Wild dogs not only make significantly more meat off of larger prey animals, but they also have a higher success catching them than they do small animals like African hares (38% success versus 31%) (see Creel & Creel, 1995 for Lycaon pictus data).  If humans were to go extinct tomorrow, dogs could never fill the role of these kinds of predators.  Simply put, dogs are more likely to try and play with a deer than to try and kill it.


Trophic diagram of organisms in relation to predators since reintroduction (Freeman et al., 2013)

As if there is very little difference, dogs are frequently labeled carnivores like their wolf cousins (implying a predatory nature); however ecological foraging models are much more nuanced than simply whether or not the food consumed is animal or plant-based.  Dogs are detritivores (i.e. scavengers—animals which live off of dead food sources).  Whether it is the kibble we drop in the bowl, the dump which feral dogs scavenge at, or even raw meat or table scraps being tossed from the table, dogs do not kill their food.  Whether feral or companion pet, the dog’s niche relies on their ability to live in close proximity to humans—a quality which is typically severely impacted by interbreeding with wolves.  Dogs utilize a very different and elongated socialization period that enables them to develop interspecies social bonds much easier (Lord, 2013), and thus the viability of hybrid offspring between dogs and wolves is severely impacted through both prezygotic and postzygotic barriers.  Quite simply, just because two animals are capable of interbreeding, claiming they are the same species does not make sense in light of almost all aspects of their phenotype outside of morphology (and even then, calling a Chihuahua a wolf is simply absurd).

Thinking about the foraging strategy of the dog as more closely related to fungi, archaea, worms, and dung beetles as opposed to the apex predator wolf might seem rather unglamorous, however in truth it highlights their ecological and evolutionary success.  All life is built on the need for energy and nutrients.  Energy needed for life comes from the sun, regardless of the species.  Plants use the energy from photons to produce sugar, which is natures way of storing energy from the sun.  For this reason, plants are termed “producers” because they create available energy and nutrients for other organisms (such as hydrogen, carbon, nitrogen, oxygen and phosphorus—among other nutrients).  Through nutrients, organisms manipulate the stored energy and use it to produce proteins that enable the organism to survive and reproduce.  The availability of these resources on a large scale is quantified in ecology as Net Primary Productivity (NPP).


Data: NASA
Image: Freeman et al., 2013


Image: Freeman et al., 2013

NPP is essentially a quantification of the available resources organisms need to survive.  If you look on the map above, you will see that NPP is highest in the tropics and lowest in the tundra.  The niche of the wolf, compared to the tropics, is in regions of the world where NPP is strikingly low—emphasizing their need to hunt and kill large prey.  Humans appropriate 24% of the NPP of the entire planet.  Think about that for a moment… nearly one-quarter of all available energy on the planet, yet we are just one of thousands if not millions of species cohabitating this blue ball in our corner of the solar system.  We can appreciate that with the laws of the conservation of energy, large consumption leads to large waste, waste that is still rich with hydrogen, carbon, nitrogen, sulfur and phosphorus.  While dogs most certainly share a common ancestor with the wolf, their emergence as a species is due to the tremendous advantage of having proximity to the largest appropriation of nutrients on the planet.  Coppinger has long emphasized the difference between “domestic” (living amongst humans) and “domesticated” (made to live amongst humans).  With no evidence that humans were ever sophisticated enough to establish breeding programs to artificially select for tame qualities like the silver fox experiment, it is not logical to believe that dogs could have emerged through careful pup selection.  Even today, it is extremely difficult to create human-socialized wolves (who still behave nothing like dogs) and inbreeding is an enormous issue within current populations—how would humans have overcome these issues when we still hadn’t become sophisticated enough to harness agriculture?


Data: Vince, 2011
Image: Freeman et al., 2013

It is hard to imagine that sometimes we forget just how much we have changed this planet.  If the consumption of nearly 25% of the planet’s NPP doesn’t make you think for a moment, then consider that 90% of all mammalian biomass on the planet consists of humans and domesticated animals.  10,000 years ago, this number was approximately 0.1%.  While rambling around since approximately 200,000 years ago, human population did not reach one billion until 1804.  By 1927, human population reached two billion.  1960: three billion.  1974: four billion.  1987: five billion.  1999: six billion.  By the year 2011, human population reached seven billion (population data and biomass percentages taken from Vince, 2011).  In parallel with increasing human population, it is estimated that there are approximately one billion dogs around the world now, whereas wolves are on the brink of extinction.  At this rate, it is only a matter of time before human population will exceed the appropriable NPP of the planet and very few undomesticated species will exist outside of detritivores feeding on human waste as the human population crashes into unsustainability.

The story is very romantic: man and wolf, hunting and foraging together.  Unfortunately there is simply no evidence; and if I’m being charitable, the probability that dogs evolved directly from grey wolves is extremely unlikely.  While many similarities are perceived to exist between dog and wolf, upon closer examination, the similarities are almost impossible to find.


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Blogging Down Our Brains

“By giving us the opinions of the uneducated, journalism keeps us in touch with the ignorance of the community.” – Oscar Wilde

Anyone interested in furthering his or her knowledge of a subject is faced with a depressing enigma: the amount of bad information outnumbers the amount of good information.  As a person who lives life trying to acquire a deeper understanding of the world, bad information really pisses me off.

Recently, a blog was published titled “5 Incredible Ways Dogs Can Read Your Mind” (Emery, 2013).  In it, the author presents several claims to support the premise that dogs can read our minds.  Formally, this is referred to as Theory of Mind, which Emery defines as the “understanding that other beings have different perceptions, and that those perceptions can be valuable” (Coren, 2011a).  This is wrong.  To be fair, even scientists do not always define it accurately, however choosing a blogger to reference who actually took the time to read the conclusions and discussions of the studies they were reviewing might have been a beneficial start.

The correct definition:

Theory of mind is the ability to make accurate inferences to understand the behavior of other animals because of abstract (theory-like) representations of the causal relationship between unobservable mental states and observable behavior (Premack & Woodruff, 1978, as cited in Penn & Povinelli, 2007—emphasis my own).

More concisely, theory of mind requires the ability to know that the behavior of another animal is a product of their cognitive state—this is distinctly different from responding to environmental factors, including that of an animal’s behavior (Udell & Wynne, 2011).  While many authors have described theory of mind as dated and potentially no longer useful (e.g. Horowitz, 2011), the goal of this blog isn’t to provide alternative evidence as to whether or not theory of mind is a valid concept in non-human animals such as dogs—best to let that war rage on in the academic community.  Instead, this blog is two-fold: 1) to correct the dirge of fallacies that went viral with Emery’s blog and 2) to use it as a model for understanding how important the source of information is.  With that as our base, let’s start at the top.  If you haven’t read the blog I am referring to yet, here is the link.

Note: the titles of each section below correlate to the paragraph titles of Emery’s blog and are here to give sign posts regarding what statements I am pulling apart: they are not claims I am making.

#5: Dogs are Capable of Empathy

“Yawning is a phenomenon directly connected to empathy, and as such has only been found to occur in species capable of empathizing (i.e. humans, and other primates), and only then within a single species.” (Emery, 2013)

First, yawning behavior is widespread and believed to be common to ALL vertebrates: including mammals, birds, fish, amphibians and reptiles (Baenninger, 1997).  Second, there are EIGHT different hypotheses regarding the function of yawning, if it is indeed even functional, and so the communication (empathy) hypothesis is just one of several ‘stabs’ at why animals yawn (Guggisberg et al. 2010).  Problematically, the communication hypothesis is not unilaterally supported and the studies that do support it are plagued with a lack of controls to rule out competing hypotheses that would be direct confounds to the results.  Even if we accepted that contagious yawning as a function of empathy was viable and true, researchers employing more stringent methods have been unable to conclude that dogs show signs of contagious yawning behavior (Harr et al., 2009).

“So obviously dogs have an uncanny ability to read our emotions … but how? Well, it’s because all humans, whether right- or left-handed, display our emotions predominantly on the right side of our faces.” (Emery, 2013)

You’d think with a 50/50 chance she might have gotten this one right merely by dumb luck—but no, humans display their emotions predominantly on their LEFT side, not their right (Borod et al. 1997).  Regardless of problems between lefts and rights, associative learning is an alternate explanation for any gaze bias observed in dogs.  However, perhaps ironically, it could even be argued that if dogs could read our mind, then they wouldn’t need to check in with the more emotional side of our face to know if we are just slightly angry about the Christmas ham getting eaten, or really angry.  Empathy is an equally hot topic as theory of mind and definitions historically have tended to overlap—often stressing the importance of “cognitive perspective taking” (Davis 1983).  Despite the sticky separation of these concepts, empathy refers to the ability of one individual to infer and share the emotional spectrums of another (Gallese, 2003; Völlm et al., 2006).  Thus, regarding whether or not non-human animals have empathy, gaze research simply cannot possibly answer the question.

#4: Dogs Understand That Your Visual Perspective Is Different from Their Own

Yes, they do, but does this constitute evidence that they possess theory of mind?  Opponents to the “perspective taking” element supporting theory of mind make a compelling counterargument: simply that all animals learn.  If a moose walks into a tree, they do not turn around and walk the other way because at some point in their life they learned they could walk around it.  A group of gazelles foraging and scanning the environment have learned to scan for predators because being eaten by a cheetah sucks.  A prey animal thus wouldn’t survive very long without some knowledge of potential threats in the environment.  Knowing this, if a gazelle stops eating, freezes, and looks across the field, the fact that all the gazelles are likely to stop foraging and check for danger does not prove the presence of theory of mind, because alternatively they could be responding due to empirical knowledge that it is in their interest to keep a look out for hungry kittens.

If you’re walking along the street and you see someone looking up, we are likely to look up as well.  The novelty of seeing someone looking up is a pretty strong stimulus to evoke our social facilitation (looking up as well), just like the gazelle and their knowledge that the environment contains dangers to be aware of, we understand that pianos or stock brokers falling on our head is also likely to put a dent in our afternoon.

Many researchers have demonstrated how dogs and wolves have varying abilities to search around visual barriers (e.g. Bräuer et al., 2004; Range et al., 2011; Virányi et al., 2009); however, ultimately here is what you have to decide for yourself:

  1. The dog is reading your mind and knows that you are looking at an object around the corner
  2. The dog notices that your eyes are looking somewhere to their right or left (an observable behavior) and is curious to investigate – oh there is a barrier?  Hmmm, let me walk around that

I think this research is interesting, but even so, this does not constitute evidence for theory of mind because it does not rule out competing hypotheses; such that a dog could be simply taking information from a visual environment—not attenuating to the cognitive states to understand the causal relationship between unobserved mental states and observed behavior.  Just my opinion, argument 2 seems much more practical and is further supported by our understanding that dogs are extremely sensitive to gazing since it is one of the most common signals they use in agonistic (conflict) behavior.

More egregiously though, Emery continues and states that dogs will abandon all morality and go for a piece of food the second you close your eyes, or turn your back, or place a barrier between you and the food, and this is a complete misinterpretation to the research done on this phenomena and thus absolute gibberish.  Leaving a food item alone is trainable.  Browse around YouTube and you will find plenty of videos where a dog is told to wait before his or her dinner bowl is set down, and then the owner walks out of the room, or even the house, before returning to release the dog to eat the food.  Honestly, a solid ‘leave it’ is one of the easiest behaviors to train, so this kind of research has to be interpreted very carefully regarding what it actually means, if anything, for the lives of our dogs.

#3 Dogs Assume That You Know Something They Don’t

As if I wasn’t already pounding my head against the desk, the author then uses the observation that dogs want to eat what we are eating as support for doggie mind reading abilities.  Unfortunately, this is not a trait unique to dogs (or humans for that matter).  Many social mammals select food preference by their group’s behavior.  For example, rats learn from group members how to determine what to eat and will learn to avoid the smell of poisoned food, a neophobic response—this is one of the reasons why rat poison doesn’t eliminate rat populations (Galef & Clark, 1971).  It is a fascinating behavior, but it does not require mind reading—rather rudimentary social facilitation.

#2 Dogs Understand Pointing

“…but the fact of the matter is that dogs and humans are the only two species currently clinging to our big blue spaceball who understand the point of pointing.”  (Emery, 2013)

Other than wolves (Udell et al., 2008), cats (Miklosi et al., 2005), parrots (Giret et al., 2009), bats (Hall et al., 2011), Jackdaws (Von Bayern & Emery, N., 2009), goats (Kaminski et al., (2005), dolphins (Pack & Herman, 2004), fur seals (Scheumann & Call, 2004), Ravens (Schloegel et al., 2007; for a review, see Udell et al., 2012)… hmm, only two species you say?  Monty Python jokes about the Spanish Inquisition aside, the ability for an animal to learn that they can walk around a tree is no different from the ability to learn that a finger might be directing towards food.  Animals who learn this distinction are socialized to people—period.  Nobody has snatched a dog that has never seen a human, tossed it in a room, pointed at a cup with food inside, and seen the dog dive in and say “thank you master!”  No, it would be shaking in the corner terrified for its life.  Animals who have been socialized to humans respond to pointing and other human communicative gestures (e.g. gazing and pointing with foot): pick your species.  Variance in this skill can be as easily explained by the failure for many animals to follow directions (just ask any school teacher how many times they have to remind students to write their name at the top of a test).


#1 Dogs Know When You Like Someone Else More

Finally, Ms. Emery claims that oxytocin is a “love- and jealousy-related hormone” (Coren, 2011b).  This claim comes from a single study involving humans playing a computer game (Shamay-Tsoory et al., 2009), however the conclusions the authors make can be reinterpreted to fit the standard functional understanding of oxytocin (Tops, 2010).  Oxytocin is a mammalian hormone that triggers milk letdown in nursing females and is involved in a wide variety of social behaviors: such as increasing pleasure during orgasm, increasing time of social contact, facilitating memory of sexual partners, protects fetal neurons from injury during delivery, improves navigational strategies, and works with vasopressin receptors to aid pair-bonding (Breedlove et al., 2010).

If you don’t see references: assume the author is an idiot

It should be clear by now that cracked.com might be just about the worst source for dog behavior science, and if you have been following some of the citation trails, Psychology Today might appear as a questionable source as well.  There are more authors writing books and blogging about dogs than there are dogs in family homes and they range from people with high school diplomas to PhDs. Citations and a reference list is an excellent way to begin to decipher the quality of information, however it is not everything either.

While bad information frustrates the daylights out of me, ultimately, the burden falls on the consumer to be sure to examine the evidence. This is one of many reasons why a list of references is so important, and why it is best to assume the author is likely an idiot if they don’t bother to acknowledge the sources of their information in a clear, concise reference section at the end.  If there is no reference list, than be sure to ask yourself whether you believe you are reading an opinion piece or an opinion piece veiled as accurate science.


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Bräuer, J., Call, J., & Tomasello, M. (2004). Visual perspective taking in dogs (Canis familiaris) in the presence of barriers. Applied Animal Behaviour Science, 88(3–4), 299–317. doi:10.1016/j.applanim.2004.03.004

Breedlove, S. M., Watson, N. V., & Rosenzweig, M. R. (2010). Biological Psychology: An Introduction to Behavioral, Cognitive, and Clinical Neuroscience, Sixth Edition (6th ed.). Sinauer Associates, Inc.

Borod, J. C., Haywood, C. S., & Koff, E. (1997). Neuropsychological aspects of facial asymmetry during emotional expression: A review of the normal adult literature. Neuropsychology Review, 7(1), 41–60.

Coren, S. (2011a). Can Your Dog Read Your Mind? Retrieved September 23, 2013, from http://www.psychologytoday.com/blog/canine-corner/201106/can-your-dog-read-your-mind

Coren, S. (2011b). Do Dogs Feel Jealousy and Envy? Retrieved September 23, 2013, from http://www.psychologytoday.com/blog/canine-corner/201111/do-dogs-feel-jealousy-and-envy

Davis, M. H. (1983). Measuring individual differences in empathy: Evidence for a multidimensional approach. Journal of Personality and Social Psychology, 44(1), 113–126. doi:10.1037/0022-3514.44.1.113

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Gallese, V. (2003). The roots of empathy: the shared manifold hypothesis and the neural basis of intersubjectivity. Psychopathology, 36(4), 171–180. doi:72786

Giret, N., Miklósi, Á., Kreutzer, M., & Bovet, D. (2008). Use of experimenter-given cues by African gray parrots (Psittacus erithacus). Animal Cognition, 12(1), 1–10. doi:10.1007/s10071-008-0163-2

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

Hall, N. J., Udell, M. A. R., Dorey, N. R., Walsh, A. L., & Wynne, C. D. L. (2011). Megachiropteran bats (Pteropus) utilize human referential stimuli to locate hidden food. Journal of comparative psychology (Washington, D.C.: 1983), 125(3), 341–346. doi:10.1037/a0023680

Harr, A. L., Gilbert, V. R., & Phillips, K. A. (2009). Do dogs (Canis familiaris) show contagious yawning? Animal Cognition, 12(6), 833–837. doi:10.1007/s10071-009-0233-0

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Kaminski, J., Riedel, J., Call, J., & Tomasello, M. (2005). Domestic goats, Capra hircus, follow gaze direction and use social cues in an object choice task. Animal Behaviour, 69(1), 11–18. doi:10.1016/j.anbehav.2004.05.008

Miklósi, Á., Pongrácz, P., Lakatos, G., Topál, J., & Csányi, V. (2005). A Comparative Study of the Use of Visual Communicative Signals in Interactions Between Dogs (Canis familiaris) and Humans and Cats (Felis catus) and Humans. Journal of Comparative Psychology, 119(2), 179–186. doi:10.1037/0735-7036.119.2.179

Pack, A. A., & Herman, L. M. (2004). Bottlenosed dolphins (Tursiops truncatus) comprehend the referent of both static and dynamic human gazing and pointing in an object-choice task. Journal of comparative psychology (Washington, D.C.: 1983), 118(2), 160–171. doi:10.1037/0735-7036.118.2.160

Penn, D. C., & Povinelli, D. J. (2007). On the lack of evidence that non-human animals possess anything remotely resembling a “theory of mind.” Philosophical Transactions of the Royal Society B: Biological Sciences, 362(1480), 731–744. doi:10.1098/rstb.2006.2023

Range, F., & Virányi, Z. (2011). Development of Gaze Following Abilities in Wolves (Canis Lupus). PLoS ONE, 6(2), e16888. doi:10.1371/journal.pone.0016888

Scheumann, M., & Call, J. (2004). The use of experimenter-given cues by South African fur seals (Arctocephalus pusillus). Animal cognition, 7(4), 224–230. doi:10.1007/s10071-004-0216-0

Schloegl, C., Kotrschal, K., & Bugnyar, T. (2008). Do common ravens (Corvus corax) rely on human or conspecific gaze cues to detect hidden food? Animal cognition, 11(2), 231–241. doi:10.1007/s10071-007-0105-4

Shamay-Tsoory, S. G., Fischer, M., Dvash, J., Harari, H., Perach-Bloom, N., & Levkovitz, Y. (2009). Intranasal Administration of Oxytocin Increases Envy and Schadenfreude (Gloating). Biological Psychiatry, 66(9), 864–870. doi:10.1016/j.biopsych.2009.06.009

Tops, M. (2010). Oxytocin: Envy or Engagement in Others? The Striatum, Psychopathy, and Molecular Mechanisms of Addiction, 67(1), e5–e6. doi:10.1016/j.biopsych.2009.08.032

Udell, M. A. R., Dorey, N. R., & Wynne, C. D. L. (2008). Wolves outperform dogs in following human social cues. Animal Behaviour, 76(6), 1767–1773. doi:10.1016/j.anbehav.2008.07.028

Udell, M., & Wynne, C. (2011). Reevaluating canine perspective-taking behavior. Learning & Behavior, 39(4), 318–323. doi:10.3758/s13420-011-0043-5

Udell, M. A., Spencer, J. M., Dorey, N. R., & Wynne, C. D. (2012). Human-socialized wolves follow diverse human gestures and they may not be alone. Int. J. Comp. Psychol, 25, 97–117.

Virányi, Z., Gácsi, M., Kubinyi, E., Topál, J., Belényi, B., Ujfalussy, D., & Miklósi, Á. (2008). Comprehension of human pointing gestures in young human-reared wolves (Canis lupus) and dogs (Canis familiaris). Animal Cognition, 11(3), 373–387. doi:10.1007/s10071-007-0127-y

Völlm, B. A., Taylor, A. N. W., Richardson, P., Corcoran, R., Stirling, J., McKie, S., … Elliott, R. (2006). Neuronal correlates of theory of mind and empathy: A functional magnetic resonance imaging study in a nonverbal task. NeuroImage, 29(1), 90–98. doi:10.1016/j.neuroimage.2005.07.022

Von Bayern, A. M. P., & Emery, N. J. (2009). Jackdaws Respond to Human Attentional States and Communicative Cues in Different Contexts. Current Biology, 19(7), 602–606. doi:10.1016/j.cub.2009.02.062

Myths, Legends, and Science: Part 2

[Part 1]

In my previous blog, I ended by poking fun of the people who superficially cling to the word “science” as the end all of reasoning, however what I did not mention is the other spectrum of people who see science as a never ending string of contradictions and thus find no value in it.  These people often dismiss the importance of science, even though they drive a car to work, fly to Cabo, write emails, take aspirin, receive vaccinations for deadly diseases, and watch movies on a phone (which is supposedly smart) that fits in their pocket.  Clearly science is just spinning its wheels.  Joking aside, these criticisms draw from the self-correcting nature of science, however this is actually a virtue if the way in which scientific theories are arrived at is fully understood.


Theory vs Hypothesis

If you could x-ray science, it could be said that its bones are made up of theories and hypotheses.  It is important to distinguish here that the colloquial use of the term “theory” is vastly different from its use in science.  Theories, much like hypotheses, make predictions, however a theory involves predictions that are rarely (if ever) unsupported because empirical tests have corroborated them numerous times.

For example, when Darwin proposed natural selection as the mechanism for evolution, he created numerous predictions that could not be readily tested in his time.  However, with the incredible advances in molecular biology, we have been able to examine variations in the sequencing of nucleic acids between organisms (i.e. DNA analysis).  Among many other things, Darwin predicted that humans and primates both evolved from a common ape-like ancestor.  Supposing that Darwin was correct, then we would expect to see fewer mutations in the DNA sequences between humans and primates then between humans and cats—since according to the theory, humans and other primates share a common mom and dad much more recently than humans and cats do.  Thus, the finding that humans share about 95% of their DNA with chimpanzees is just one of many findings that corroborates the theory of evolution.1


Credit: M.F. Bonnan

Other theories you may be familiar with are atomic theory, the theory of relativity, germ theory, cell theory, and so on and so forth.  Often, people will try to impugn a theory by saying statements to the effect of, “evolution is just a theory.”  However, once someone understands how theories are established, this statement is as ridiculous as saying “germs and atoms are just theories.”  Really what they mean to try and say—although falsely—is that they are hypotheses, as if to suggest that science wasn’t really sure one way or the other.

Hypotheses come in many varieties, they can be based on lots of evidence, some evidence, or sometimes no evidence at all, and the longer they bounce around without being refuted, the stronger the evidence becomes.  Atomic theory, for example, started off as a hypothesis back with the pre-Socratics around 400ish BCE.  Naturally, scientific exploration has fine-tuned it like a small European sports car, however the initial hypothesis has managed to maintain its central idea: that the universe is composed of atoms.  Evidence supporting atomic theory can be seen in particle accelerators, when you toss salt in water, cook oil in a pan, or even when you watch ice melt into water.

Hypotheses become theories when they are extensively supported with experimental and observational data

kurt-lewin-good theoryKurt Lewin (1952)2


The philosopher Karl Popper emblazoned the importance of falsifiability in science, and while many of Popper’s ideas were tremendously controversial, the importance of falsifiability in science is one that everyone seems to happily agree on.

I found that those of my friends who were admirers of Marx, Freud and Adler were impressed by a number of points common to these theories and especially by their apparent explanatory power.  These theories appeared to be able to explain practically everything that happened within the fields to which they referred.  The study of any of them seemed to have the effect of an intellectual conversion or revelation, opening your eyes to a new truth hidden from those not yet initiated.  Once your eyes were thus opened you saw confirming instances everywhere: The world was full of verifications of the theory.  Whatever happened always confirmed it.  Thus its truth appeared manifest; and unbelievers were clearly people who did not want to see the manifest truth, who refused to see it, either because it was against their class interest or because of their repression which were still “unanalysed” and crying aloud for treatment.  (Popper, 1963)


Einstein provides a very nice example of how falsifiability plays a role in science even when a theory seems extremely intangible.  In 1905, Einstein declared in his publication of the Special Theory of Relativity that E=MC2.  In 1916, Einstein then published the General Theory of Relativity, which predicted that light bends with the distortions of space and time (i.e. is affected by gravity).  The claim was so radical—so audacious—that the astrophysicist and philosopher, Arthur Eddington, made an expedition to an island on the western coast of Africa to test Einstein’s claim.  If Einstein was correct then the stars near the Sun would be in a different position than normally anticipated because the distortion of space around the Sun would have caused the light to bend—and vice versa, if the stars near the Sun did not change position, then the theory could be rejected.  This kind of testability is implicit in determining whether a hypothesis or theory is falsifiable.  As you might have guessed, Eddington’s photographs of the stars near the Sun during the solar eclipse successfully demonstrated Einstein’s predictions.  Initially, if I had asked you to think of a test where you could determine whether or not light bends, you would probably scratch your head for a while (as would I).


Photograph from the island of Principe

It is a great exercise to think about the testability of a claim by thinking about what evidence would be needed in order to falsify it.  For example, Freud claimed that conflicts between an individual’s conscious and unconscious mind resulted in neurotic behavior; problematically, since by definition the unconscious mind is “not available to introspection,” it is not testable.  If it is not testable, it is not falsifiable.  If it is not falsifiable, it isn’t science.

Assumptions: the pitfall of a good hypothesis

Naturally, the results of experiments do not always support the hypothesis, however this does not mean that the hypothesis is wrong.  The magic of good critical thinking is finding where a hypothesis might be carrying hidden baggage—more specifically: assumptions.  Remember Semmelweis and the use of chlorinated lime solution?  Semmelweis hypothesized that if the cause of childbed fever was in fact a contamination of putrid matter from the morgue, then the introduction of something that would kill the putrid matter (i.e. an antiseptic) would result in stopping the contamination.  While he guessed well, this hypothesis carried an assumption that a chlorinated lime solution works as an antiseptic.  However, since this predates germ theory, nobody even knew what an antiseptic was (let alone an effective one).  An even stronger example of the difficulty with assumptions played out historically in what is called Stellar Parallax.

A great debate that began in ancient Greece and did not end until Copernicus in the 16th century was whether the universe is Geocentric (Earth at the center) or Heliocentric (Sun at the center).  Being clever at geometry, the Greeks thought of a way to test the two competing hypotheses.  If the universe was heliocentric, then there would be a calculable difference between the stars in the sky at one time of the year versus another—however, if the universe was geocentric, then there would be no change.  The Greeks measured, and sure enough, they concluded that no, the stars did not seem to change, ergo the universe must be geocentric.

You can replicate this test by closing your left eye and extending your thumb out in front of you, pick an object you can cover with the width of your thumb (ideally about 3-10ft away), then open your left eye and close your right (i.e. switch the closed eye)—alternating back and forth you should see the object(s) behind your thumb bounce back and forth.  Looking at the diagram below, “July” would be your left eye’s perspective and “January” would be your right eye’s perspective.


Stellar Parallax

The first problem for the Greeks was that they never imagined that the stars were over 40 trillion kilometers away (4 light years), thus they didn’t realize how small the changes they were looking for really were—for perspective, the Earth is 40,000km in circumference, so you would have to complete a million journeys around the Earth a million times in order to cover the distance between us and our next closest star Alpha Centauri. The second problem for the ancients was that they didn’t know which star to try and pick out to use as a reference point, so it was a hypothesis with assumptions that were nearly impossible to know at the time.

The moral of the story is that just because a test fails a hypothesis, we cannot simply throw it away because the negative result might simply be the outcome of an uncontrolled confound in the experimental design.  This means that if the results of an experiment do not support a hypothesis, we go back and think, “what were my controls” “do I have any assumptions in my methods” “are there confounds I haven’t controlled for” etc.

The Scientific Method

Putting all these pieces together, you will probably recall from one of your high school science classes a flow chart that looks somewhat like this:


Scientific Method – Version 1.0

I call this Version 1.0 because this is the scientific method in a vacuum.  Science historian Steven Shapin, a professor at Harvard University, makes a compelling argument that one of the most important elements which shaped the Scientific Revolution was the way in which scientists began working together to coordinate research.3  This emphasizes an aspect of the scientific method that version 1.0 is completely lacking, and that is its unique social structure.


Scientific Method – Version 2.0

Science cannot evolve if the knowledge of the past is too unreliable to expand on, nor can it evolve without the influence of society and the benefits it has on the human condition.  Peer-review is a function required for the highest level of publication and capitalizes on how scientists maintain correspondence and quality control; thus, while the quality can vary, peer-review and editorial standards are one of the most significant elements of good science. They are the measure by which all science is scrutinized for miscalculations, methodological flaws, invalid conclusions, and sometimes, deception.

Image Credits:

Scientific Method 1.0 – (http://courses.washington.edu/esrm430/sm.jpg)

Scientific Method 2.0 – (http://arstechnica.com/science/2009/03/building-a-better-way-of-understanding-science/)


(1) Britten, R.J. 2002. ‘Divergence between samples of chimpanzee and human DNA sequences is 5% counting indels.’ Proceedings National Academy Science 99:13633-13635

(2) Lewin, K. (1952). Field theory in social science: Selected theoretical papers by Kurt Lewin. London: Tavistock.

(3) Shapin, S. (1996).  The Scientific Revolution.  Chicago: University of Chicago Press.

Myths, Legends, and Science: Part 1

One of the most common words in today’s dog industry is the word “science”: such as science-based, scientifically proven, or backed by science.  Problematically, the word is often utilized in an attempt to punctuate that an idea, product, method, or concept is simply fact, and that anyone who disagrees with it is likely ignorant, uneducated, or just plain wrong.  Sure, arguments and debates are essential to science, and in order for those to happen we have to have strong opinions.  However, there are a growing number of people who have started resorting to the word “science” without knowing the methods or conclusions that constitutes the evidence behind their claims and with the extra assumption that science only has a singular opinion beyond reproach.  The problem is that this isn’t really what science is about, nor is it how we got to where we are today.


What is Science?

Most people would agree that physics, chemistry, and biology are science—often referred to as the “natural” sciences as they passionately try to unravel the mysteries of the universe.  Although what about other subjects, such as mathematics?  Music?  Astronomy?  Philosophy?  Psychology?  Economics?  Metaphysics?  Logic?  What makes one subject a science in our minds and another a pseudo-science?  While my goal is not to simply list what subjects I personally believe are and are not science, by the end of my series I hope you can make that determination for yourself, it is interesting that as far as history is concerned, ‘science’ is actually a relatively new word.  The roots of what is now modern (western) science began in ancient Greece with the advancements of philosophy and the infamous Aristotle (384 to 322 BCE).

As a philosopher, Aristotle’s celebrity was incalculably immense.  Described in Dante’s Inferno as “the master of those who know,” Aristotle wrote about the world and the heavens in ways that still permeate modern science.  His greatness as a philosopher set the stage so strongly that for about 2,000 years, those we would call scientists throughout history actually referred to themselves as “natural philosophers.”  This can be seen as late as even Isaac Newton’s publication of Philosophiae naturalis principia mathematica (Mathematical Principles of Natural Philosophy) in the 18th century.


Aristotle and Zeno’s Paradoxes

Zeno of Elea was a philosopher from the 5th century (BCE)—although what a great name for an evil space villain.  Zeno was an extreme rationalist who argued that it was reason, and reason alone, that could give us the gateway into an understanding of the way things are.  He believed that the senses (i.e. seeing, hearing, smelling, touching) were tainted as a tool for building knowledge and used several paradoxes as evidence that even observations about motion (as in an object moving) were actually just figments of the senses.

Zeno’s Dichotomy Paradox (dichotomy literally means “cutting in two”): imagine a dog running towards a stationary object.  The object is at a finite distance D and the running happens in a finite time of T.  Zeno claimed that in order to travel D, the dog must first travel the first half of D, then half of the distance that remains, followed by half the distance that remains of that, followed by half the distance that remains of that as well (i.e. the dog would travel half of D, then a quarter of D, then an eighth of D, then a sixteenth of D, then 1/32nd of D, 1/64th of D…) etc. ad infinitum—infinitely.  Following this logic, it would then have to be assumed that the dog will have to travel an infinite number of distances in a finite amount of time.  To Zeno, this was a contradiction; therefore, assuming that an object moves because we see it move is a false assumption when it is simply illogical for an object to complete an infinite number of distances within a finite amount of time.


Aristotle, however, came along with a resolution to Zeno’s Dichotomy Paradox.  Instead of arguing with the conclusion, even though clearly the conclusion is absurd, Aristotle created a resolution to the paradox by focusing on the assumptions within the argument.  By focusing on the paradox’s construction, Aristotle demonstrated two of the most important elements that science is built on—reasoning and logic—which is what made him so infamous (not his primitive hypotheses of the heavens which is often what is taught in history class).  While Aristotle’s resolutions are apt, the paradox wasn’t laid to rest until modern mathematics came up with a mathematical proof to rationally explain Zeno’s Dichotomy.1

ImageModern notation for solving Zeno’s Paradox

Deductive vs Inductive Reasoning 

There are two distinct forms of reasoning that can be used to make a claim: deductive and inductive reasoning.  Deductive reasoning takes a very large premise and narrows it down to a smaller conclusion.

  1. All dogs have noses.
  2. Muffy is a dog.
  3. Therefore, Muffy has a nose.

The power of deductive reasoning is that when the premises are true, and the argument construction is valid, the conclusion is undeniably true—I don’t know many people who would argue that Muffy doesn’t have a nose.  However, at the same time, deductive reasoning can be tricky because it could be built on a false premise.  Here is another deductive argument:

  1. All atoms have one or more protons.
  2. Carbon is an atom.
  3. Therefore, carbon has one or more protons.

We can only say that this is undeniably true (i.e. “sound”) if we have examined every atom in the universe.  However, despite not having examined every atom in the universe, we still accept that all atoms have one or more protons because of Inductive Reasoning.  Inductive reasoning would look like this:

  1. Every atom we have found so far has one or more protons.
  2. Therefore the next atom we find will have one or more protons.

If we assume this is true, based on this inductive reasoning, that the next atom we find will have one or more protons, then it is sound to conclude that carbon has one or more protons.  However, inductive reasoning cannot prove because it generalizes from a finite sample, thus it is able to suggest that a hypothesis is probably true.  Like a car with no warranty: it makes no guarantees.

Semmelweis and ‘Childbed Fever’

Ignaz Semmelweis was a Hungarian physician whose story expands this concept of the importance of inductive reasoning in science.  In the mid-19th century, Semmelweis worked at a hospital in Vienna where there were two maternity divisions, however problematically, about 12% to 17% of the women who entered the First Division to give birth began to subsequently die with what was called childbed fever (a horrific death with symptoms including organ failure and edema), while only about 2-3% of the woman who entered the Second Division suffered the same fate.  Systematically, Semmelweis formed several hypotheses to try and discover the cause of the mortality rate in the first division.

The first hypothesis was that the deaths were due to Atmospheric Influences.  Before germ theory, people believed epidemics were passed through atmospheric events, however to Semmelweis, this seemed impossible because it did not explain why women who gave birth on the street on route to the hospital had a higher survival rate than the women in the first division, nor why two different wings of the same hospital would consistently have different mortality rates—so this hypothesis was thrown out.  Other hypotheses included: overcrowding; giving birth on the back instead of the side (it was common for women to give birth on their sides at this time); diet; rough handling by doctors; and Death by the terrifying and debilitating presence of Priests (my personal favorite even though it was unsupported)2.  While many of these were also thrown out due to a lack of logic or probability, Semmelweis ran experiments where he had the priests take different routes through the hospital and where he had all the mothers in first division give birth on their sides instead of on their back—no luck.

After almost four years of trying to solve the problem, a colleague of Semmelweis’ received a puncture wound from a student’s scalpel in the morgue and died of the exact same symptoms as the women of first division.  It suddenly occurred to him that the medical students—who not coincidentally had begun additional training by performing autopsies on cadavers about four years prior—were often traveling straight from the morgue to the delivery room and often still smelled of rotting flesh (believe it or not, medicine really has come a long way).  Semmelweis then instituted a protocol that medical students had to wash their hands in a chlorine and lime solution before heading to the delivery room in the first division.  Because this predates germ theory, Semmelweis had only decided to try a chlorinated lime solution because it was effective at removing the smell accumulated from working on cadavers.  Regardless, within no time at all, the mortality rate in the first division dropped 90%.  Sadly however, mandatory hand washing caused a huge uproar and Semmelweis was politically ruined (despite having evidential vindication—i.e. women stopped dropping like flies) for even suggesting that invisible putrid matter derived from dead and living organisms might be the cause of the mortality rates.  Eventually, despite his discovery, Semmelweis was dismissed from the Vienna hospital only to then be forced to move back to Budapest due to harassment from the Vienna medical community before eventually being committed to a mental institution.  Apparently doctors really didn’t want to have to wash their hands…


The first vertical line represents the beginning of autopsy investigation in Vienna (Wien)
The second vertical line represents the introduction of handwashing procedures

Important to the question of science, however, is that even though Semmelweis solved the problem it turned out that his hypothesis was actually still somewhat incorrect.  As it turns out, the women dying from childbed fever were actually dying from a genial tract sepsis often caused by bacterial infections of Staphylococcus (staph infections)—not putrid matter derived from living and cadaverous organisms.  One could argue, “well, what’s the difference?”  If I suggested (as Einstein did, and emphasized by my incredible high school physics teacher) that gravity did not involve gravitons but rather bends and distortions in space and time, you would agree they are two significantly different hypotheses, regardless of the observational outcome.  Inductive reasoning is a powerful tool, however conceptually and historically we know that it creates an understanding about probability, not fact.

Science is…

Dictionary: Science is the intellectual and practical activity encompassing the systematic study of the structure and behavior of the physical and natural world through observation and experiment.

ImageBroken down, we get two parts.  The first part starts with the idea of the intellectual and practical nature of science.  The history of science is filled with the Galileans (those who, like Galileo Galilei, believe in science for the sake of science—the intellectual nature) and the Baconians (referring to those who, like Francis Bacon, believe science has to have a purpose—the practical nature).  More broadly and simply we can summarize this as the two primary types of scientific investigation: theoretical and applied.  The theory of relativity would be exemplary of theoretical science and research in medicine would be exemplary of applied science.

The second part of the definition very generally describes what is known as the “scientific method” which I will go into more detail in my next blog.  For now it is suffice to understand that thanks to great thinkers like Aristotle, science is built on experience (i.e. it is empirical).  The method of science utilizes techniques designed to solve conceptual problems of our experience in the real world.  Why is the sky blue?  Why do dogs like to hump certain people’s legs?  Why does coffee wake us up?

While this definition is a great start, science is also much more.  For instance, science is falsifiable; it is exploratory; it is beholden to concise and logical arguments; it is damaged by bias; and most importantly, it is ever changing.  Science does not ascertain facts, nor does it establish truths.  Science is about examining the current evidence, asking new questions, and modifying our preexisting conclusions based on new explorations.

So the next time someone uses the word “science” as a definitive proof for an argument, remember that a true scientist is both cautious and careful when making claims and would never stoop so low as to insult the intelligence of someone by defaulting to the word “science” to win an argument against them—especially without establishing who’s experiences they are referring to.

Blog Continued in Part 2



(1) An awarded and readable overview of Greek science and philosophy can be found in G.E.R Lloyd, Greek Science After Aristotle (New York, NY: W.W. Norton & Company, Inc., 1973)

(2) For more information about Semmelweis and his life, including detailed accounts and translations of his writing, check out W.J. Sinclair, Semmelweis: His Life and His Doctrine (Manchester, England: Manchester University Press, 1909)

Image sources

Science proves you’re wrong: Zazzle.com
Newton’s Principae Naturalis Mathematica: NPR.org
Zeno’s Dichotomy Paradox: http://berto-meister.blogspot.com/2013/04/what-is-zenos-dichotomy-paradox.html
Notation for sum of an infinite series: Wikipedia
Table of mortality data from Vienna hospital: Wikipedia
Warning Science in Progress: Zazzle.com
“Science” image: http://www.gdfalksen.com/post/52184550214

Pseudo-behavioral science

Recently I saw a study from the Journal of Experimental Analysis of Behavior shared via the American Veterinary Society of Animal Behavior (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1397788/#!po=50.0000).  In it, the authors claim that a higher rate of reinforcement for a behavior creates a stronger resistance to the extinction of the behavior when reinforcement is removed: a very broad claim given the niche experiment.  Reading the abstract, most anyone would be happy to accept their claim, especially professionals who are always on the prowl for more evidence to support their particular belief system.  However, this is a great example of why we have to be careful about what sources we decide constitute science.

There are several problems in the JEAB study linked above:

First, two experiments with 3 to 4 starving rats (of one species) in strict confinement cannot be expected to explain the behavior of other healthy animals such as dogs:

“The subjects were 4 male Long Evans hooded rats, about a year old at the start of the experiment. Obtained as juveniles (about 150 g), they were gradually (over several months) brought to a weight of 335 g (± 15 g) and maintained at that level by free access to food blocks in their home cages for 1 to 1.5 hrs after each session. (Ator, 1991, provides a rationale for this method of food deprivation for rats)” [emphasis my own].

It should be highlighted that one of the rats died after condition 6 and a second rat did not follow one of the extinction conditions because it appeared ill.  Yet the deprivation, which resulted in illness or death in 50% of their animals, is rationalized and considered necessary.

Second, it is unclear if they actually found anything.  In addition to the small population and no statistically significant findings, this study is a general discussion on mathematical principles, not behavioral observations.  Both experiments reported in the study required manipulation of their data in order for it to fit their hypothesis.  Let me repeat, the authors willingly admitted to throwing out data they ‘didn’t like’.  The authors justify this as removing an outlier, but some pause has to be taken because it is not scientific to willfully remove data in order to prove a hypothesis or theory.  Thankfully, the authors do contribute this paragraph appropriately:

“Basing conclusions solely on adjusted data, however, can be risky. For any set of data, some adjustment can be found to generate whatever new relation one might wish for. If the adjustment is selected arbitrarily, the relation that emerges will be arbitrary as well and thus misleading about the relevant behavioral processes. The question, then, is whether a particular adjustment can be justified on grounds beyond its ability to produce a particular outcome.”

It is beyond the realm of my understanding that radical behaviorists believe that this formula accurately depicts the phenomenon of the process of behavioral extinction, regardless of species, function, and ecology: log(Bx/B0) = -x(c+dr)/ra .  It is especially incredible to me that such hypotheses are being generated due to results that are undergoing fraudulent statistical p-hacking whereby statistics are calculated over and over and populations and data adjusted until the authors find the results they are looking for (which in this study couldn’t even result in any results being statistically significant).  A real scientist would never throw out a chunk of their data so they could prove a mathematical formula fit a complex biological behavior, nor would they observe the death and illness of half their animals as something only needing mention in a footnote of the appendix.

This is not science; this is torture and mathematical perversion.

An interview with Marc Bekoff

Marc and Bessie

Marc and Bessie

Few names are as synonymous with “pioneer” as Marc Bekoff. An authority on behavior, sociality, and play in mammals (first through canids and then more broadly), Dr. Bekoff has a truly large and wide-ranging bibliography to match his incredible career. He has brought us hundreds of peer-reviewed scientific research publications, essays, and book chapters, and has written 23 books on animal emotions, cognition, play, and compassionate conservation. In 2000, he co-founded the organization Ethologists for the Ethical Treatment of Animals with Jane Goodall. Among his numerous honors, he was presented with the Exemplar Award from the Animal Behavior Society for his major long-term contributions to the field of animal behavior and the St. Francis of Assisi Award by the Auckland (New Zealand) SPCA. Ethologist and ethicist Marc Bekoff has been an extraordinary advocate for the welfare of all animals and it is one of my greatest honors to interview him here today.

What inspired you to study animal behavior?

Bekoff: I was always interested in animal behavior. My parents told me that when I was around 3 years old I began ‘minding animals’ and always asked them what animals were thinking and feeling. I published a book called ‘Minding Animals’ in 2002 based on this conversation with my wonderful parents. I love learning about other animals and still write about them in scientific and mass-market books, and essays for Psychology Today.

Marc Bekoff and Adam Miklosi with wolf

Marc Bekoff and Adam Miklosi with wolf

Why is play so much fun?

Bekoff:  Play is fun because when animals including humans are playing they are relaxed and stress free, and simply are able to enjoy themselves with their family and friends. Play is also contagious—just watch dogs join their buddies at a dog park and frolic on and on and on. This always make me smile and want to join in.

How important is play when trying to understand the behavior of social mammals?

Bekoff:  Play is incredibly important because it is essential that animals play when they are young in order to become socialized, card-carrying members of their species, and also for them to get physical activity and cognitive training. Two colleagues and I have suggested that play is ‘training for the unexpected’.

Have the ethics of behavioral research changed over the course of your career?

Bekoff:  Absolutely. It seems like every week something ‘good’ is happening for further protecting animals from wanton and horrific abuse in a wide variety of venues. Nonetheless, there is still a lot of work that needs to be done to protect animals from being abused.

What role has research played in discovering the inner lives of animals, including dogs, and how does this affect research and how we should treat animals in the future?

Bekoff:  Rigorous scientific research has clearly shown that other animals are sentient beings, very intelligent, extremely emotional, and moral, and that we must never cause intentional and unnecessary pain, suffering, and death.

Marc with Cormorant

Marc with Cormorant

If I’m not mistaken, Michael W. Fox was a teacher for you in grad school. What was it like being a student of Michael’s?

Bekoff:  Yes indeed he was. He was an exemplary mentor and always was there for me. I learned an incredible amount from Michael in many different arenas, lessons that I tried to use with my own students. Michael was always a forward-looking thinker and walked his talk. He was and remains an inspiration to me and to many others.

Your schedule of speaking engagements is remarkable, what brought you to SPARCS?

Bekoff:  I like very much bringing what we know about dogs and other animals to a wide audience because the fields of animal behavior and cognitive ethology, the study of animal minds and what’s in them, are incredibly exciting. Almost daily we’re learning more and more about the amazing cognitive and emotional capacities of other animals and it’s essential to share this information with as wide an audience as possible.

Would you say that your relationship with dogs and other canids have shaped aspects of how you view the world? If so, in what way?

Bekoff:  Yes it has. I’ve learned many life lessons about trust, friendship, devotion, kindness, compassion, empathy, and love from the animals with whom I’ve shared my home, the land around my home, and from those who I’ve had the opportunity and pleasure to study. My life would have been and would continue to be empty without the nonhuman animals with whom I’ve had contact in a wide variety of situations.

Is there anything else you’d like to mention to our readers about you or the upcoming conference in Redmond?

Bekoff:  Try to attend! It’ll be a wonderful gathering and there will be a lot of information that’ll be shared with you all.

Dr. Bekoff will be speaking at SPARCS in Redmond, WA on June 28th and June 29th to discuss the emotional lives of animals and how research and science can “rewild” our hearts.

For more information regarding SPARCS, please visit www.CanineScience.info; to learn about Dr. Bekoff’s other upcoming appearances around the world and his publications, please visit his homepage at www.MarcBekoff.com. Thank you Dr. Bekoff.

Zeke likes his ball

Zeke likes his ball


This interview was originally published March 25th here

Stress and learning

The Yerkes-Dodson Law

In 1908, Yerkes and Dodson published findings of a remarkable phenomenon they discovered regarding the relationship between arousal and performance.  The law asserts:

  1. The speed of learning and performance increases with arousal; however, it quickly reaches an optimal intensity where learning and performance then deteriorate.
  2. Both weak stimuli and strong stimuli result in slow habit-formation.
  3. The more difficult the task, the lower the optimal level of arousal for maximum performance.
  4. “A stimulus whose strength is nearer to the [minimum] threshold than to the point of harmful stimulation is most favorable to the acquisition of a habit.”  (Yerkes & Dodson, 1908)

Yerkes-Dodson Law

Hebb's graph of the Yerkes-Dodson Law (image source: wikipedia)

Hebb’s graph of the Yerkes-Dodson Law (image source: wikipedia)

Two things eventually arise from dog training methods that involve prolonged arousal or stress.  First, as cognitive function deteriorates, behavior becomes resistant to extinction.  Thus if our objective is to make a particular behavior diminish or stop, in actuality the stress initially preserves the ineffectual behavior (Schwabe & Wolf, 2011).  As stress continues to move in the “high” range of the graph above, corticosteroids are actively being pumped by the hypothalamo-pituitary-adrenal (HPA) axis to support sympathetic nervous system activity (commonly referred to as “fight or flight”).  When these systems are stimulated excessively—because the physical well-being of an animal is at harm—an animal’s learning for a survival strategy impedes learning of the task/behavior (Joels et al., 2006).  Eventually, as learning and performance come to a halt, a dog enters a state of learned helplessness.

Harsh punishment and the “Hang” – a dog’s fight for survival

In a case study published in the Journal of Veterinary Behavior (Grohmann et al., 2013), a German Shepherd was diagnosed with severe cerebral edema resulting in ischemic brain damage from a training session with its owner.  When the dog failed to perform desirably, the owner “disciplined” the dog by hanging it on its leash with an equipped choke chain for 60s.  Brain ischemia, or cerebral ischemia, is caused when there is a lack of oxygen rich blood flow to the brain causing cerebral hypoxia and brain death—in layman’s terms: death or brain damage by strangulation.  Other presenting conditions included:

  • Anxiety
  • Panting
  • Tachycardic (racing heart beat)
  • Severe disorientation
  • Pleurothotonus (abnormal and sustained involuntary muscle contractions)
  • Blindness
  • Bilateral mydriasis
  • Left-sided facial motor paralysis

Medical differentials for the presenting conditions included diffuse axonal injury, vascular ischemia, increase in intracranial pressure, and hemorrhage.  The dog was euthanized after the diagnosis was made and the owner declined a postmortem examination.

All I can think about when I read stories like this is: what was going through the mind of that poor dog, an emotional and sentient being, as the choke chain cut off blood and oxygen to the brain?

What is dog training

Dog training is literally the act of teaching a dog to perform a certain task—whether it is a sit or to stop lunging on leash—a task which ultimately requires the dog to make a successful decision.  This is a complex physiological process that involves 3 things: the cognitive processing of relevant information, the estimation of relationships between actions and their potential consequences, and use of executive functions to optimize decision-making performance (Mair et al., 2011).

Prolonged and severe stress has deleterious effects on cognitive function, memory formation, and performance (McEwen & Sapolsky, 1995; Yerkes & Dodson, 1908)

Owners and trainers need to abandon confrontational methods and techniques, teach dogs ‘English as a Second Language’, talk to them like a loving parent, learn their cognitive landscape by playing and interacting with them, and enjoy the fact that they are not a robot: they are an individual with complex biological emotions and thoughts.


Grohmann, K., Dickomeit, M. J., Schmidt, M. J., & Kramer, M. (2013). Severe brain damage after punitive training technique with a choke chain collar in a German shepherd dog. Journal of Veterinary Behavior: Clinical Applications and Research. doi:10.1016/j.jveb.2013.01.002

Joëls, M., Pu, Z., Wiegert, O., Oitzl, M. S., & Krugers, H. J. (2006). Learning under stress: how does it work? Trends in Cognitive Sciences, 10(4), 152–158. doi:10.1016/j.tics.2006.02.002

Mair, R. G., Onos, K. D., & Hembrook, J. R. (2011). Cognitive Activation by Central Thalamic Stimulation: The Yerkes-Dodson Law Revisited. Dose-Response, 9(3), 313–331. doi:10.2203/dose-response.10-017.Mair

McEwen, B. S., & Sapolsky, R. M. (1995). Stress and cognitive function. Current opinion in neurobiology, 5(2), 205–216.

Schwabe, L., & Wolf, O. T. (2011). Stress increases behavioral resistance to extinction. Psychoneuroendocrinology, 36(9), 1287–1293. doi:10.1016/j.psyneuen.2011.02.002

Yerkes, R., & Dodson, J. (1908). The relation of strength of stimulus to rapidity of habit-formation. Journal of Comparative Neurology and Psychology, 18, 459–482.

A conversation with Michael W. Fox

A love for animals

A love for animals

Michael W. Fox, who is not only a veterinarian with a PhD degree in medicine but also holds a Doctor of Science degree in ethology/animal behavior from the University of London, England, shows how science can enhance our empathy and ability to better care for and communicate with animals, and speak for the rights and interests of all creatures great and small.  An Honor Roll Member of the American Veterinary Medical Association and a founding member of the International Association for Applied Animal Ethology, Dr. Fox has helped make animal behavior and animal welfare science part of the veterinary teaching curriculum, and more generally awakening public sensibility through his internationally syndicated newspaper column Animal Doctor both in the U.S and in many other countries.  As an advocate of holistic and integrative veterinary medicine, and as a long term member of the American Holistic Veterinary Medical Association, he has promoted advances in organic agriculture, pet food nutritional quality, and is part of an international community of scientists, bioethicists and others opposing the incorporation of genetically engineered animals and crops into the human and animal food-chain by the non-sustainable industrial agricultural food and drug industrial complex.

Michael and I chatted for about 40 minutes, and it was such a moving conversation that I thought I would provide his voice to accompany his words in hopes they inspire you as they have me.

[Fox on the field of Ethology]

Fox:  You break down “Ethology” and it’s [two parts]: -ology is the study of, like “biology” or “cosmology”, and Eth-ology is the study of ethos, or the spirit that moves animals.  You are essentially studying the spirit that is motivating this being to behave, to respond, to interact.  I remember the conference in Rennes several years ago that was opened in the town hall by Konrad Lorenz—I mention it in one of my books at least—where he said to the gathering of these international ethologists, “before you are going to study an animal, you are going to have to first love it.”  I was standing with Daniel Lehrman and some of the other more mechanistic ethologists from the United States and they all started saying, “ohhh, he’s going soft” and “not being scientific”, but what Konrad was saying is that you have to have the connection—which I call empathy—before you can really begin to understand what’s going on.  And that was easy for him because he studied the Greylag geese and they imprinted onto him when he raised them as infants.  As fresh little hatchlings they thought he was their mother so they imprinted onto him and it was very easy.  But he did say that some animals can be very difficult to love, and he had one graduate student who was set studying some little aquatic creature who spent most of the day on the bottom of the aquarium tank and just came up for a gulp of air and then went down to the bottom and he said, “some animals are difficult to love but you have to hang in there.”

But I would say, getting on the practical level with dog trainers, broken down from the need to have an objective Skinnerian pigeon in the conditioning box to simply being with the animal is a big step for that individual to take.  And then you find yourself in the space with the animal and free from your own limiting conceptual space and that’s a real letting go, which is the antithesis of control and obedience training.  That letting go you enter another dimension really, of a deeper rapport and a communion where you are dealing not only with cognitive processes but emotional things going on too: such as fear, shyness, the need for attention, all those motivating factors that John Paul Scott and John Fuller, my mentors in Bar Harbor, looked at, and Daniel Freedman in his studies of different ways of raising different breeds of dogs, and genetics and emotions play a big role in trainability, but ultimately when you can break through the emotional and genetic related differences in temperament, dogs do turn out to be very similar in their basic intelligence (that was the conclusion of Scott and Fuller in their work).

[Fox on his early research in epigenetics and development]

Fox:  In my early research, I was interested in what is called “epigenetics” and my doctoral dissertation which was published by the University of Chicago Press, “Brain and Behavioral Development in the Dog”, I looked at other researchers who had been doing early handling studies on mice, Victor Denenberg and Gig Levine in particular, who found that if they gently handled the mice when they were pregnant, the offspring were more docile and more exploratory and they learned better because it affected their cognitive processes and emotionality and adrenal pituitary axis or whatever else was going on.  And so I applied these principles to some puppies that I raised and some that those that I had a programmed early life history of handling doing various things to them, including mild stress but lots of loving and cuddling, that they behaved in far more emotionally stable ways and were quicker learners than litter mates that were not handled the same way, and we applied that to the army’s German Shepherds that were being raised for combat work during the Vietnam war and we developed the Super Dog program based on that early research and these dogs performed incredibly better than dogs who hadn’t been in the program in field and they saved a lot of lives.

The reason I had the opportunity to speak with Michael is because he is speaking this summer at the first annual SPARCS International Conference on Dog Behavior in Redmond, WA.  Given the demand for Michael’s incredible knowledge and ideas, I had to ask him: why SPARCS?

[Fox about his speaking engagement at SPARCS]

Fox:  I think the opportunity to engage with the public, especially those who have dogs, especially those who are concerned about animals, and especially those who are in any professional realm as dog handlers, trainers and professional care givers including [veterinary technicians] and veterinarians, that I still think I have something to say that is old and eternally new: so far as socialization, stages of development, how early experiences can affect later behavior, the animal’s immune system and how modern interventions in the animals lives from vaccinations to using these various so-called flea and tick preventive drugs to manufactured foods (including GMOs) [that] are affecting dogs and cats, and my incredible information input that I get from my syndicated newspaper column of the kinds of behavioral and health problems that we’re seeing today, and how that spectrum has changed over the 30, 40 years that I’ve been writing this nationally syndicated column.  I have a lot of new stuff to share.

Michael will be presenting on all three days at SPARCS this summer, from June 28th-June 30th.  For more information on SPARCS you can visit CanineScience.info.  For research documentation and verification of the virtues of enlightened self-interest in humanity adopting Dr Albert Schweitzer’s advocacy of Reverence for all Life, visit Dr. Fox’s website www.drfoxvet.com and www.Facebook.com/DrFoxvet