DOMESTIC ANIMAL BEHAVIOR
Animal behavior relates to what an animal does and why it does it. The types of behaviors
exhibited are rich and various. Some are genetically determined, or instinctive, while others are
learned behaviors.
Definition of animal behavior :
Animal behaviour is the expression of an effort to adapt or adjust to different internal and
external conditions, i.e. behaviour can be described as an animal’s response to a stimulus. Animal
behavior is the scientific study of everything animals do, regardless of whether the animals are
single-celled organisms, invertebrates, fish, amphibians, reptiles, birds, or mammals. It involves
investigating the relationship between animals and their physical environment as well as to other
organisms, and includes such topics as how animals find and defend resources, avoid predators,
choose mates and reproduce, and care for their young.
Prehistory and an Adaptive Perspective on Behavioral Observations
The behavior of animals evolves and is shaped by
natural selection. In a similar way our own
behaviors, our understanding of how animals
behave, was shaped by survival needs in the
remote past. By better understanding the behaviors
of animals, our hunter-gatherer ancestors more
successfully caught and trapped game. There is of
course no way to see direct evidence of such
observational skills in prehistoric humans as they
are no longer in existence. However, samples of
Paleolithic art from 40,000+ years ago provide
indirect evidence that primitive humans observed the animals that they hunted. Cave paintings
often portray animals that are naturally found in herds with like members of their species. Images
seem to capture the mass movement. These paintings show hyenas hunting in groups. Bears are
portrayed as solitary. In some cases the solitary animals show up together, but they appear to
interact and face each other in some contest.
By studying animal behavior primitive humans were able to exploit the differences in behaviors
associated with solitary animals versus those living in herds. Their knowledge helped them
capture prey. They learned that animals traveling in herds could be driven over cliffs in large
numbers provided that lead animals were first driven over the precipice. Some aspects of human
behavior and hunting were cultural transmitted. They learned to avoid risky situations where
large predatory beasts could ambush them, or they died. Some human behavior with survival
value became instinctive. In a curious way, our own initial ideas regarding behavior undoubtedly
developed for the very reasons behaviors have arisen in all organisms -- behavior has adaptive
value and is shaped by the force of natural selection. Even the process of learning behavior was
shaped by natural selection.
Although the meaning of cave art is debatable, it is clear that human's appreciation of animals
reaches back to the dawn of prehistory. In modern day, if one is ever granted the opportunity to
follow an aboriginal "tracker", one can learn an amazing amount about an animals behavior from
just a few signs in the sand. A student of animal behavior uses similar skills of observation when
they study their organism of choice. Many field biologists become extraordinary "trackers"
because they must catch many animals repeatedly over the years. Many of these animals become
incredibly difficult to catch as the animals themselves learn to predict the researcher's behavior.
One might say that we are very adapted for the study of animal behavior owing to the force of
past natural selection.
Despite this kind of intuitive sense of animal behavior, it is still a large leap the practical aspect of
behavioral observations of animals to the study of animal behavior as a discipline. What are the
origins of modern ideas on animal behavior? The scientific study of animal behavior is founded
on Darwin's ideas concerning evolution by the process of natural selection (Darwin, 1859). In
treating the ideas in any field, one must consider the origin of those ideas. This appreciation of
philosophy is essential for complete comprehension of important concepts. We could use
Darwin's theory of evolution by the process of natural selection as a starting point for modern
ideas on animal behavior, but realize that our understanding of animal behavior has very deep
roots indeed and undoubtedly arose during our own prehistory.
Instinctive Behaviors
Web making by spiders is an example of a genetically determined or instinctive behavior. There
is little variation between individuals in how they construct the web and it is constructed similarly
each time they do it. Ethologists (people who study behavior) call such a behavior a fixed action
pattern. Fixed action patterns do not require learning or prior experience for their expression.
They can, however, be very complex. For example it has been noted that a cocoon-spinning
spider performs over 6000 individual movements in a virtually identical fashion each time it
prepares and closes its cocoon.
Fixed action patterns cannot be identified solely because they are highly stereotypic and species-
specific. For example, songs of bird species fit those criteria but are actually learned behaviors,
not instinctive behaviors. Birds deprived of the experience of hearing the song do not produce the
characteristic song pattern. Deprivation experiments in which animals are raised without
parents,or contact with their own species, have typically been used to help distinguish a behavior
that is a fixed action pattern from those which are learned.
The lack of expression of a behavior in a deprivation study does not necessarily mean that it is
not genetically determined. It may mean that the requisite stimuli are absent. Stimuli to elicit the
expression of fixed action patterns are called releasers. Also a fixed action pattern may not be
displayed because the animal is not in the appropriate physiological or developmental state. For
example courtship behaviors are not shown by pre-pubertal animals even if the appropriate
releasers are present.
Learned behaviors.
Learning is the modification of behavior in response to specific experiences. Learned behaviors
of animals can be classified in various ways:
In Associative Learning an animal learns to associate one stimulus with another. There are 2
forms of associative learning. The first is described as classical conditioning and was
demonstrated by Pavlov in his famous dog experiments. First he stimulated dogs to salivate by
rubbing meat powder on their lips. The meat powder odor stimulated salivation as a physiological
process. He then "conditioned " dogs by ringing a bell, or a tuning fork, at the same time as
applying the meat powder. He then demonstrated that the animals had become conditioned to
associate the sound of the bell with the meat powder and would salivate to the noise without food
present. A second form of associative learning is called operant conditioning. In operant
conditioning an animal conducts a chance action e.g. pressing a lever and is rewarded with food.
Rapidly the animal learns that the action leads to a food reward and will carry out the behavior
repeatedly for food. This is the type of approach applied in most animal training.
Observational learning or modelling is when the animal learns a behavior through watching
other animals conduct the behavior. For example,in a pack animal such as the wolf, hunting
behaviors, fit this category.
Insight learning is in a sense the "highest form" of learning observed. It is the ability to problem
solve or to perform a correct or appropriate behavior the first time the animal is exposed to a
situation. For example a chimpanzee may stack boxes to obtain a food object hung out of its
reach without ever having seen this solution to the problem before. However, it is not restricted to
primates e.g. Ravens and other birds will also show insight learning.
An animal may cease to carry out a response to a stimulus if the appropariet response no longer
occurs. For example using the operant conditioning example given above if pressing the lever no
longer leads to a food reward the behavior will probably become less frequent and stop.
Nature versus Nurture.
A debate among human psychologists, as well as animal behaviorists, concerns the relative
importance of instinctive and learned behaviors. The factor influencing learned behaviors is the
environment in which the animal is placed. This debate is often referred to as Nature (genes)
versus nurture (environment).
The balance between fixed and learned behaviors varies with species. In humans, a large part of
our behaviors are learned. In the absence of role models to learn from (deprivation) there is a
greater emphasis on programmed behaviors. Some behaviors need to be fixed for survival
because there is unlikely to be a second chance to learn them. For example the kangaroo rat
instinctively reacts to the sound of a rattlesnake by executing an escape jump. This is a species
specific defense response.
With learned behaviors there ar often critical or sensitive
periods for the development of the appropriate learned
behavior. For example a dog that has not been socialized
to humans by 14 weeks of age is unlikely to be a good
pet. Dogs that have not been well socialized to other dogs
may be frightened of them or will not breed with them.
Imprinting is another example of a process that must
occur within a distinct, usually short, time period. It is
also irreversible and involves an attachment to an object
that will evoke subsequentadult behaviors and can be
generalized to all examples of the object.
The classic example is the work of Lorenz with goslings in which they imprinted on Lorenz. The
imprinted adult geese directed courtship behavior to Lorenz rather than other geese.
Konrad Lorenz. Nobel Laureate
Animal Cognition
Cognition is the ability to think. When applied in the context of animal behavior it refers to the
ability of an animal to be aware of and make judgements about its environment. Are animals
conscious of themselves and their place in the world? Do they feel pain, pleasure and sadness in
the same way as humans? Are the differences between animals and humans in cognitive ability
absolute, as believed by Descartes, or a matter of degree? Professor Donald Griffin of Princeton
has been a proponent of the concept of cognitive ethology. This views conscious thinking as an
inherent and essential part of the behavior of many non-human animals. Conscious thinking has
been demonstrated in non-human primates but it is a very difficult subject to study with scientific
rigor. As we have discussed above very many complex behaviors are actually genetically
determined or learned by simple associations that do not need cognition. Difficult though they are
to study the answers to questions about animal cognition may profoundly affect our views on the
treatment of animals and their welfare.
Follow the link to an article on animal thinking by Pam Noble entitled the Status of the Animal
Mind
Communication
Communication among animals relies upon their abilities to perceive sensory information. This
may be visual, auditory or olfactory. Domestic animals perceive the world in a different fashion
to us so may respond differently also.
Vision - An obvious example of seeing the world differently relates to eye placement. Horses are
set laterally, providing a wide range of monocular vision (215 degrees) but a small 60-70 degree
arc of binocular vision. In contrast, the cat has a much smaller arc of vision (around 180 degree)
because of the placement of the eyes to the front but more than half of this is binocular vision.
When the relative visual acuity of domestic animals is compared, the ranking is pig, sheep, cattle,
dogs and horses. All the domestic animals have some ability to discriminate colors and most have
superior night vision to humans.
Auditory sensing - Again there are differing ranges of hearing, as well as different acuities. Sheep
and dogs can discriminate higher frequencies than humans. Cats range of hearing is similar to
humans.
Olfactory senses - This is perhaps the most important sense of domestic animals in terms of
communication. Dogs probably have the greatest olfactory ability of the domestic species. They
are able to detect many compounds at 1/100 the concentration of humans and for many weeks
after they were placed. Odors and pheremones are very important triggers for domestic animal
behavior.
The communication methods employed by an animal in response to stimuli perceived through
these sense organs are the same types: auditory, visual (facial expressions, posture, grooming) ,
and olfactory. These communication methods have varying importance between domestic
species:
Auditory - Pigs have perhaps the most complex set of domestic animal vocal sounds - more than
20 distinct types have been identified. Horses, cats and dogs also have many sounds but those of
cattle and sheep have been little studied. The categories of calls include greeting, distress,
separation, excitement, aggression, fear, pain and so on.
Visual - In the horse, for example, the ear position can tell a lot about the animals disposition.
Ears pointed back generally denote aggression and the flatter the ears to the head the greater the
aggression. Another visual expression in male horses and ruminants is the Flehmen response.
This is a curl of the lip when the urine of a female in estrus is smelled. In the cat, a high tail is a
greeting or sign of being curious. Raised hackles, and lips drawn back in the dog denotes
aggression; if the ears are flat against the head and the tail is between the legs this signifies fear.
Pigs and horses show grooming behavior, as do many of the monkeys and primates. Subordinate
pigs groom dominant ones. Horses tend to groom horses of comparable rank. Cattle and cats
spend long periods of time self-grooming but little time grooming others. Subordinate cattle will
lick dominant cattle.
Olfactory - Olfactory clues and scent serve to mark territory , show the way home, and
distinguish individuals. Urine and feces are powerful means of olfactory communication. Cats
and dogs also have anal sacs which are additional scent organs. Olfactory stimuli play a major
part in heat detection in cattle, and for distinguishing animals. The ewe will accept an orphaned
lamb if it is made to smell like her own lamb.
Animal Behavior and the Animal Scientist.
Domestic animals display the same ranges of behaviors that would be seen in other animals and
have both instinctive and learned behaviors. Understanding domestic animal behavior is not only
fascinating and intellectually stimulating it is also crucial in their management, productivity and
welfare.
Behavior and the detection of animal disease.
Often the experienced veterinarian or owner who has a sound understanding of normal animal
behavior can identify the existence of a problem from a change in posture, sounds etc. Some are
very easy to recognize such as acute laminitis others can be quite subtle. An animal kicking at its
belly probably has an abdominal pain, while sweating in horses is a sign of acute pain e.g. colic.
Dogs with ear infections often tilt their heads. Excessive grooming or scratching is indicative of
parasites or irritations. A change in the order in which an animal comes into the milking parlor,
particularly if a dominant cows hangs back, may indicate illness.
Control of Breeding and Feeding
The definition of a domestic animal is one which has been altered by selective breeding and
control of the food supply by humans. An understanding of behavior allows the detection of
animals in heat, can be used to identify feeding problems, modify maternal behavior to accept
orphaned animals, manipulate group numbers and size of animals to reduce aggresion and so on.
Training
In training domestic animals heavy reliance is placed upon associative conditioning i.e., positive
reinforcement of desired behaviors, but the genetic blueprint is also very important. For example
the genetic predisposition of the border collie to herd is adapted by consistent training to increase
skills and control of that instinctive ability. This is a particularly crucial aspect of animal science
with respect to the companion and recreation species. The use learning theory is also applied to
help animals relearn to reduce vices and bad habits.
Behavior and the Environment
Observation of animal behavior provides a great deal of information about the extent to which the
animal copes with the environment in which they are kept. It can also contribute to the design of
better housing and management systems for farm animals. An obvious example is climate.
Temperatures which are either too hot or too cold will lead to behavioral changes which are
readily detectable. If dairy cows are standing in free stalls rather than feeding or lying down it
probably indicates that the stalls are uncomfortable. If the environment of pigs is very barren and
unstimulating this will lead to some aberrant behaviors such as aggresion and also to stereotypes.
This is discussed further under the animal welfare section.
Bibliography
Craig, James. (1981). Domestic Animal behavior: Causes and Implications for Animal Care and
Management. Prentice-Hall;, Inc. Englewood Cliffes, New Jersey
Houpt, Katherine. (1991) Domestic Animal Behavior for Veterinarians and Animal Scientists.
Iowa State Press, Ames, Iowa,
The Traditions of Animal Behavior: Nature versus Nurture
Ethology
Although Darwin shifted the way we view animal behavior, the discipline also has a tradition that
stretches before the time of Darwin (Drickamer and Vessey, 1986). The field of ethology, which
is the study of the evolution and functional significance of behavior, originated with C. O.
Whitman in the 1800's. Whitman coined the term instinct to describe the display patterns of
pigeons. The ethogram, a graph of the time course or switch points in a sequence of behaviors,
became a way of categorizing species-typical behaviors. Many of these instincts are triggered by
various environmental stimuli and von Uexkull termed such triggers of instinctive stereotyped
behaviors sign stimuli. A classic stimulus triggers the courtship display of male three-spined
sticklebacks fish. The enlarged belly of a female triggers the zig-zag dance in male stickleback
fish. The males use the dance to entice the female stickleback to enter the nest that the male has
built.
Much of the work of early ethologists was synthesized by two Nobel Laureates, Niko Tinbergen
and Konrad Lorenz. Lorenz is noted for his work on genetically programmed behaviors in young
and for studies on imprinting, during critical developmental periods in young. A classic example
of imprinting occurs in young geese when they form an image of parent just after hatching. If the
hatchlings first encounter a human such as Lorenz, they will imprint on him and follow him
around as if he were their mother. A third Nobel Laureate, Karl von Frisch, pioneered studies in
bee communication and foraging.
One of Tinbergen's seminal contributions to Behavior was to formulate a method studying animal
behavior (Tinbergen, 1963). This method forms the basis for how I have structured material in
this text. These issues are central to developing a philosophical approach to animal behavior. The
ethological approach had a strong Darwinian tradition underlying its development. Much of the
work in ethology was aimed at understanding the ultimate evolutionary reasons for behavior.
Tinbergen listed four areas of inquiry that could be used to understand issues of animal behavior.
The following mnemonic can be used to remember these four areas ABCDEF [Lehrman, 1965]:
A -- Animal refers to the organisms.
B -- Behavior refers to the observable actions of the organism.
C -- Causation refers to the proximate causes of behavior such as genes,
hormones, and nerve impulses that control the expression of behaviors.
D -- Development refers to the ontogeny of behaviors such as imprinting, or in
the case of cognition, learning.
E -- Evolution refers to the phylogenetic context in which behaviors are found.
For example, the prevalence of parental care in birds, but not reptiles (with some
exceptions) is an example of the taxonomic affiliations of some behaviors.
F -- Function refers to the adaptive value or contribution that the behavior makes
to fitness.
Psychology and Behaviorism
The ethological approach typified by the research of Lorenz, Tinbergen, and von Frisch was
largely concerned with the behavior of organisms as it is expressed in their natural environment.
Another large group of scientists focussed on the mechanistic underpinnings of behavior. This
research was on model organisms (e.g., Norway rat) in a controlled laboratory setting. Classic
work by B. F. Skinner lead to the development of the use of learning paradigms, and the Skinner
Box remains an important tool in the field of animal psychology.
Figure 3. A rat learns to press a bar in a Skinner Box.
With each bar press the rat is rewarded with food.
Learning theorists sought the similarities mechanisms in
all animals that allow animals to respond to their
environment. The broadly defined field of comparative
psychology included many developments in the
psychological sciences and spanned the following topical
areas:
1. perceptual psychology -- reception of environmental stimuli through the senses, and
subjective perceptual interpretation of these sensory stimuli,
2. physiological psychology -- an attempt to relate physiological properties within an
organism to external behaviors (e.g., measuring nerve impulse transmission in sensory
and motor nerves),
3. functionalism -- the study of the mind (e.g., John Dewey) and how the mind operates.
4. behaviorism -- the study of how accumulated experiences shape the behavior of the
organism. The idea that an organisms is born a tabula rasa or (blank slate) upon which
experiences accumulate and shape behavior is central to behaviorism.
5. animal psychology -- while initially related to the study of learning in model systems,
the field of animal psychology in the present day encompasses a large body of work
related to cognition in a diverse group of animals.
The Debate on Nature versus Nurture
The field of Ethology typified by the work of Tinbergen, Lorenz, and von Frisch, and the broadly
defined field of comparative psychology formed two drastically different schools of thought on
the causes of behavior. We can compare and contrast their views to develop a deeper
understanding behavioral analysis. The field of ethology, which originated in Europe, looked to
the genetic underpinnings of behavior. In contrast the field of comparative psychology, which
originated in America, viewed behaviors as largely the product of the environment. Differences
between the ethology and animal psychology led to a debate on the causes of behavior that has
been captured in the often-quoted phrase "nature versus nature". What influences behavior --
genes or environment? The answer to this contentious debate cannot be put in terms of either
genes or the environment, but must instead be looked at in terms of a more complex interaction
between genes and the environment.
Behavioral Ecology and Sociobiology
Students of Behavioral Ecology have attempted to synthesize both the evolutionary traditions of
Ethology, and the mechanistic studies of Comparative Psychology. This is a relatively new
movement compared to the traditions of ethology and psychology and has developed over the last
three decades. The study of behavioral ecology looks at how organisms interact in their natural
environments (Krebs and Davies, 1987). Researchers are interested in both the mechanistic
underpinnings of behavior, as well as the fitness consequences of behavioral traits. This tradition
can be traced back to Tinbergen and the four study areas (Causation, Development, Evolution and
Function). Behavioral ecology is more broad than just a study of behavior, but also draws in
issues of energetics and physiology (e.g., Calow, 1987). Rather than measure differences in
survival and reproduction of behavioral traits, behavioral ecologists often use behavioral traits
that maximize energy acquisition or foraging success as proxies for fitness traits. The
development of optimal foraging during the 70's and 80's has added a distinct theoretical
perspective to the field of Behavioral Ecology.
The newest approach to studying behavior involves a consideration of social systems in a diverse
group of organisms. This field has taken off since the publication of Sociobiology by E. O.
Wilson (1980). Because some of these ideas have been applied to humans, the theory has been the
target of much controversy. Sociobiology has a strong Darwinian tradition as it attempts to
develop rules that explain the evolution of social systems and as such it.
More recently, the field of Evolutionary Psychology have co-opted the approaches of behavioral
ecology and sociobiology in order to explain a diversity of human behaviors such as foraging,
siblicide, and female choice. Humans are considered subject to the same "organic rules" that
shape other organisms. Needless to say, this area is ripe for debate as researchers attempt to
derive explanations for behaviors displayed by humans in modern society.
Ultimate versus Proximate Causes
The dichotomy between Ethology and Comparative Psychology with their concerns for
adaptation and mechanism respectively, can be succinctly described as a concern for ultimate
versus proximate causes. Ernst Mayr (1961) describe the pursuit of those ultimate causes as a
concern for the "Why Questions." Why does a bird give parental care? Why is a bee brightly
colored? In contrast, the pursuit of proximate causes is concerned with the way the world works
or the "How Questions." How does a bat transmit echoes? How do nerves carry impulses? Where
are memories stored?
Tinbergen's four study areas also block out into ultimate versus proximate causation. For
example, Tinbergen's view of causation is concerned with Proximate Causation, or mechanism.
Development is also considered to be in the category of proximate cause. However, evolution or
phylogenetic context is squarely in the field of ultimate cause, as is the issue of function as such
issues of adaptive value or fitness are directly related to evolution and evolutionary change (Curio
1994). Our study of animal behavior begins with a consideration of the ultimate causes of
evolutionary change -- adaptation and natural selection.
Cause, Development, Evolution, and Function
Tinbergen's breakdown can be used as a summary of the material covered thus far. I prefer to
make the breakdown a little more detailed to include other approaches that have been added more
recently by Behavioral Ecologists and Sociobiologists: Genes, Ecology, Physiology,
Development and Learning, Evolution, and Sociality. This categorization is slightly finer than
Tinbergen's but it provides the structure for this text and a schema for understanding the process
of adaptation in behaviors at a variety of temporal scales. Paul Sherman (1988) would add yet
another category to the list -- Cognition. However, as cognitive theory is an outgrowth of
development and learning, it will be included in those categories. Behavioral Ecology is
undergoing a large-scale renaissance as researchers attempt to generalize the classically-
developed ideas of Psychology and Cognitive Processes into wild populations (Real, 1994).
The first two subjects in the sequence Genes and Ecology will cover the basics of Darwinian
natural and sexual selection as they apply to animal behavior. To cope with environmental
variation, the organism evolves adaptations of physiology that promote successful survival or
reproduction. Such physiological changes could act at the level of endocrinology,
neurophysiology, metabolism, or any of the myriad of proximate mechanisms that operate in an
organism. These proximate mechanisms are used to help the organism cope with both abiotic
(e.g., the extremes of weather, navigation, etc.) and biotic environmental factors (e.g., the social
environment, predation, etc.). Additional components to an organism's life are the developmental
changes and learning that occur from ovum/sperm to maturity that are also adaptations to a
particular way of life. Whereas physiology operates in the very short term, development unfolds
during the lifespan of an organism. With an understanding of these genetic, ecological,
physiological, developmental and cognitive processes in hand, we will be ready to tackle the
concepts of behavioral evolution.
Phylogeny and Constraints on the Evolution of Behavior
Up to this point, I have operated under the premise that adaptation is the sole process that governs
the evolution of behavior. However, in recent years, students of animal behavior have become
more sensitive to the limitations of organic systems to change in an evolutionary sense.
Organisms may be well adapted, but limitations in organismal design constrain adaptation. In
addition, organisms are also constrained by the effects of history or their own phylogeny. During
the evolution of a lineage, adaptations pile on top of one another. The net result is that closely
related organisms share similar features which further constrain the acquisition of new
adaptations Functional and structural constraints arise from the material properties of organisms
and additional development constraints arise from how structures are built during embryogenesis.
The constraints on organisms reside at the level of proximate causation.
Consider a simple phylogenetic example taken from two lineages of vertebrates -- birds and
mammals. All birds lay eggs, undoubtedly because the common ancestor of birds, some reptile-
like dinosaur, also laid eggs. However, most mammals bear live young because in the remote past
a new kind of mammal-like reptile evolved a different mode of life and passed this novel trait on
to all subsequent species in the lineage or phylogeny. A famous exception to this mammalian
generalization includes the monotreme mammals of Australia, the platypus and echidna. It is
thought that the monotremes branched off from the main stock of mammals so early in the past
that they retain the more ancestral mode of egg-laying reproduction.
Such differences in reproductive mode (egg-laying versus live-bearing) constrain both birds and
mammals in terms of parental care behaviors that evolve in each group. Additional adaptations in
mammals may similarly constrain the evolution of parental care. Evolution of the mammary
gland as the primary source of nutrition tends to lead to species of mammals displaying a
preponderance of maternal care. There are in fact far fewer examples of male care in mammals
compared to birds. In contrast, many bird species have evolved male and female parental care
behaviors so that rearing the young can be accomplished by both parents. Some species of birds
provide a milky substance which is secreted by part of their digestive system called the crop.
Because both male and female birds have the crop, in theory both parents can evolve to produce a
milky substance as a form of parental investment. The phylogenetic difference in the amount of
male versus female care between mammals and birds leads to additional differences in how
mating systems evolve in these two groups. In order to understand phylogenetic constraints that
operate on other traits, we need a working knowledge of the proximate mechanisms, as well as
the process of natural selection. Accordingly, I leave the discussion of such higher order
macroevolutionary process for later chapters.
Societal and Cultural Evolution
Finally, I leave the discussion of sociality until the very end, because it includes even more
complex interactions that occur between organisms such as communication. The added
complexity of sociality makes the study of behavior very rich indeed. A simple example will
suffice. In developing our paradigm for animal behavior, I have thus far assumed that all changes
that are passed on between generations are largely genetic and that populations evolve and genes
change by the process of natural and sexual selection.
Social evolution and the advent of culture introduces another mode of long-standing
transmission of behavioral traits between generations. One need only walk into the nearest library
to realize the impact of mass storage of human culture has on cultural transmission of culture.
Libraries are a vehicle whereby information is passed on to subsequent generations of humans,
but there is no genetic basis to the information in libraries. The theory of cultural evolution holds
that many behavioral changes in humans might have a largely non-genetic component arising
from such cultural transmission of information. Your reading of this book forms a kind of
cultural inheritance.
Genetic Influences in Animal Behavior
Clearly, genes significantly influence animal behavior. This is the only reasonable conclusion in
cases where animals born and reared in isolation nevertheless develop age-appropriate, species-
specific behaviors that they could not have possibly learned from other individuals. Such instincts
that occur even in isolated animals include insect mating behavior; courtship, nesting, and brood-
rearing behavior of pigeons; the songs of some (not all) birds; bird flight; and nut-cracking and
nut-burying by squirrels. Animals are born "knowing" how to do certain things. Experiments that
rule out social learning and trial-and-error learning leave heredity—that is, genes—as the only
logical explanation for such behaviors.
People have long used artificial selection to produce animal breeds with desirable behavioral
traits, such as dogs that herd sheep or hunt. ArtificialExperiments with honeybees have confirmed
the relationship between genes and [Link] can also shape the reactions of fruit flies
(Drosophila) to light and gravity and the ability of rats to learn mazes. Such results would not be
possible if genes did not influence behavior. The fact that X rays and chemicals can induce
mutations that alter behavior strengthens the link to genetics. Mutations have changed obstacle-
avoidance behavior of Paramecium and biological clocks and several behaviors in Drosophila.
Experimental Evidence of Gene-Behavior Links
Breeding experiments confirm the relationship between genes and behaviors. For example,
worker honeybees normally react to diseased or dead pupae by uncapping the honeycomb cell
containing the pupa, dragging the pupa out, and removing it from the hive. This helps to prevent
the spread of infections through the colony. Experiments in which normal honeybees were
crossed with bees that do not bring out their dead traced the behavior to two genes: one that
induces workers to uncap the diseased cell, and the other that induces the insects to remove the
diseased pupa.
Hybrids between behaviorally different strains and species of animals exhibit behaviors
intermediate between those of the parents, or combine the parental behaviors. This has been seen
for aggression in honeybees, courtship in Drosophila, breeding behaviors of cichlid fishes, food
preferences in garter snakes, bird migratory and nesting behaviors, and bird distress calls.
PAVLOV, IVAN PETROVICH (1849–1936)
Russian biologist who won the 1904 Nobel Prize in physiology for his demonstration of the idea
of a "conditioned reflex." Pavlov trained dogs to drool at the sound of a bell by feeding them
immediately after sounding the bell.
LORENZ, KONRAD (1903–1989)
Austrian biologist who founded the study of animal behavior, or ethology. Lorenz said that
animal behavior evolves in the same way as physical structures, such as wings. Lorenz shared the
Nobel Prize in physiology with fellow ethologists Karl von Frisch and Nikolaas Tinbergen.
Finally, there is no such thing as a gene for any behavior. There is no aggression gene, no gay
gene, no gene for bird song or nut-burying. Genes encode proteins, nothing more; but through
proteins, they can influence behavior. Aggression and sexual behavior, for example, are
influenced by testosterone, and testosterone is synthesized by enzymes, which are proteins
encoded by deoxyribonucleic acid (DNA). Thus one can see how genes would influence these
behaviors. All behavior, furthermore, depends on chemical signals (neurotransmitters) that are
released by one neuron and bind to receptors on the next neuron. Neurotransmitters, too, are
synthesized by enzymes encoded by DNA, and their receptors are proteins as well.
Neurotransmitter levels control mood and probably aspects of personality. The list goes on and
on. Indeed, it is impossible to see how genes could not play a role in behavior.
