Ethology, a sub-topic of zoology, is the study of animal behavior that focuses on behavior in natural settings, as opposed to behaviourism, which focuses on the behavior of animals in laboratory settings. The term “ethology” is based off the Greek word ethos, which means character. It was first made popular in 1902 by William Morton Wheeler, an American myrmecologist, but the term was actually suggested by John Stuart Mill in 1843 for use in associationistic psychology. Ethology can also be associated with comparative psychology, although this is a sub-topic of psychology and bases animal behavior off human psychology.

Before the nineteenth century, the predominant thought regarding animals and humans upheld by scientists was scala naturae, created by Aristotle. This theory suggested that animals and humans were classified on a pyramid, with humans at the top and animals placed from top to bottom depending upon their complexity. In the Western world, it was thought that animals were created for specific purposes and that they were immutable. These prevailing ideas changed when Jean-Baptiste Lamarck introduced a theory of evolution that upheld two main points. This theory suggested that the organs and behaviors of animals can develop and change over time and that the changes could be passed down through generations. This theory also suggested that animals and humans alike were seeking higher levels of perfection.

When Charles Darwin visited the Galapagos Islands, he was aware of Lamarck’s theories and took inspiration from them for his own theory about evolution. Because of his book The Expression of the Emotions in Man and Animals, which many ethologists were inspired by, many consider Darwin to be the first Modern ethologist. Under the guidance of Darwin, George Romanes used a controlled method known as anecdotal cognitivism to study animal behavior and its relation to the theory of evolution, but this method was not supported in scientific world. Some ethologists, like Julian Huxley and Oskar Heinroth, focused on the natural or instinctual behaviors of animals by using an ethogram, which included the description of a common behavior of a species and the frequency at which that behavior occurred in a particular, natural situation. This data could be updated and checked by other ethologists, creating a collective base of information.

It is thought that Konrad Lorenz, under the tutelage of Oskar Heinroth, identified fixed action patterns, which were used to identify animal communication techniques. The most in-depth study using this method was conducted by Karl von Frisch, who studied bee communication. Niko Tinbergen, a colleague of Lorenz, posed four points for ethological studies. These include the function of a behavior, the causation of the behavior, the development of the behavior, and the possible evolutionary history behind the behavior.

Because of the advancements made by Tinbergen and Lorenz, ethology expanded in Europe until World War II. After this, Tinbergen moved to the UK where he began working at  University of Oxford, helping to expand ethology in that area alongside other ethologsts like Robert Hinde and William Thorpe. Ethology was also becoming popular in North America during this time.

Ethology focuses on many topics including instinct, relying heavily on the idea of fixed action patterns.  Tinbergen worked with these kinds of studies using fish, butterflies, and birds to prove that outside influence of instinctual factors, like breeding preferences, could be manipulated while still revealing the natural behaviors of the animals. Learning, which comprises many techniques including habituation, imprinting, associative learning, and imitation, is also an important focus in ethology. Habituation is displayed when an animal disassociates a behavior with a stimulus, such as squirrels ignoring the alarm calls of other squirrels that often give false warnings.

Associative learning is displayed when an animal reacts to a stimulus that can or cannot be related to another stimulus, like a goldfish rising to the surface of the water before it is fed. Other focal points of ethology include social structure and mating, both of which can hold complicated stimuli and behaviors.

In 1970, John H. Crook published a pivotal paper suggesting that most of the work done in ethology prior to this time was comparative and that more focus would need to be given to the social aspects of groups of animals, rather than single individuals.  In that same year, Robert Ardrey published a book called The Social Contract: A Personal Inquiry into the Evolutionary Sources of Order and Disorder, which focused on the comparisons of animal and human behavior. More recently, ethology has expanded to include sociobiology, traditional comparative psychology, and animal cognition, among other areas of study. Many businesses now utilize ethology including animal psychology, anthropology, and veterinary practices.

Image Caption: This is a collection of images intended to show a diversity of animal behaviour (ethology). Credit: DrChrissy/Wikipedia (CC BY-SA 3.0)