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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 animals 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. t 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. !rehistory and an Adaptive !erspective on "ehavioral #bservations The behavior of animals evolves and is shaped by natural selection. n a similar way our own behaviors, our understanding of how animals behave, was shaped by survival needs in the remote past. "y 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 s$ills in prehistoric humans as they are no longer in existence. %owever, samples of !aleolithic art from &','''( years ago provide indirect evidence that primitive humans observed the animals that they hunted. )ave paintings often portray animals that are naturally found in herds with li$e members of their species. mages seem to capture the mass movement. These paintings show hyenas hunting in groups. "ears are portrayed as solitary. n some cases the solitary animals show up together, but they appear to interact and face each other in some contest. "y studying animal behavior primitive humans were able to exploit the differences in behaviors associated with solitary animals versus those living in herds. Their $nowledge helped them capture prey. They learned that animals traveling in herds could be driven over cliffs in large numbers provided that lead animals were first driven over the precipice. Some aspects of human behavior and hunting were cultural transmitted. They learned to avoid ris$y situations where large predatory beasts could ambush them, or they died. Some human behavior with survival value became instinctive. n 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. *ven 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 bac$ to the dawn of prehistory. n modern day, if one is ever granted the opportunity to follow an aboriginal ,trac$er,, one can learn an ama-ing amount about an animals behavior from just a few signs in the sand. A student of animal behavior uses similar s$ills of observation when

they study their organism of choice. .any field biologists become extraordinary ,trac$ers, because they must catch many animals repeatedly over the years. .any of these animals become incredibly difficult to catch as the animals themselves learn to predict the researcher+s behavior. #ne might say that we are very adapted for the study of animal behavior owing to the force of past natural selection. /espite this $ind 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. 0hat are the origins of modern ideas on animal behavior1 The scientific study of animal behavior is founded on /arwin+s ideas concerning evolution by the process of natural selection 2/arwin, 34567. n 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. 0e could use /arwin+s theory of evolution by the process of natural selection as a starting point for modern ideas on animal behavior, but reali-e that our understanding of animal behavior has very deep roots indeed and undoubtedly arose during our own prehistory.

Instin tive Behaviors 0eb ma$ing 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. Etholo!ists 2people who study behavior7 call such a behavior a fi"e# a tion $attern. 8ixed action patterns do not re9uire learning or prior experience for their expression. They can, however, be very complex. 8or example it has been noted that a cocoon-spinning spider performs over :''' individual movements in a virtually identical fashion each time it prepares and closes its cocoon. 8ixed action patterns cannot be identified solely because they are highly stereotypic and speciesspecific. 8or example, songs of bird species fit those criteria but are actually learned behaviors, not instinctive behaviors. "irds deprived of the experience of hearing the song do not produce the characteristic song pattern. /eprivation experiments in which animals are raised without parents,or contact with their own species, have typically been used to help distinguish a behavior that is a fixed action pattern from those which are learned. The lac$ of expression of a behavior in a deprivation study does not necessarily mean that it is not genetically determined. t may mean that the re9uisite 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. 8or example courtship behaviors are not shown by pre-pubertal animals even if the appropriate releasers are present. Learne# behaviors% Learnin! is the modification of behavior in response to specific experiences. ;earned behaviors of animals can be classified in various ways< n Asso iative Learnin! an animal learns to associate one stimulus with another. There are = forms of associative learning. The first is described as lassi al on#itionin! and was demonstrated by !avlov in his famous dog experiments. 8irst he stimulated dogs to salivate by

rubbing meat powder on their lips. The meat powder odor stimulated salivation as a physiological process. %e then ,conditioned , dogs by ringing a bell, or a tuning for$, at the same time as applying the meat powder. %e then demonstrated that the animals had become conditioned to associate the sound of the bell with the meat powder and would salivate to the noise without food present. A second form of associative learning is called o$erant on#itionin!. n operant conditioning an animal conducts a chance action e.g. pressing a lever and is rewarded with food. >apidly 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 learnin! or mo#ellin! is when the animal learns a behavior through watching other animals conduct the behavior. 8or example,in a pac$ animal such as the wolf, hunting behaviors, fit this category. Insi!ht learnin! is in a sense the ,highest form, of learning observed. t is the ability to problem solve or to perform a correct or appropriate behavior the first time the animal is exposed to a situation. 8or example a chimpan-ee may stac$ boxes to obtain a food object hung out of its reach without ever having seen this solution to the problem before. %owever, it is not restricted to primates e.g. >avens 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. 8or 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 fre9uent and stop. Nat&re vers&s N&rt&re% 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 ?ature 2genes7 versus nurture 2environment7. The balance between fixed and learned behaviors varies with species. n humans, a large part of our behaviors are learned. n the absence of role models to learn from 2deprivation7 there is a greater emphasis on programmed behaviors. Some behaviors need to be fixed for survival because there is unli$ely to be a second chance to learn them. 8or example the $angaroo rat instinctively reacts to the sound of a rattlesna$e by executing an escape jump. This is a species specific defense response. 0ith learned behaviors there ar often critical or sensitive periods for the development of the appropriate learned behavior. 8or example a dog that has not been sociali-ed to humans by 3& wee$s of age is unli$ely to be a good pet. /ogs that have not been well sociali-ed to other dogs may be frightened of them or will not breed with them. Im$rintin! is another example of a process that must occur within a distinct, usually short, time period. t is also irreversible and involves an attachment to an object that will evo$e subse9uentadult behaviors and can be generali-ed to all examples of the object.

The classic example is the wor$ of ;oren- with goslings in which they imprinted on ;oren-. The imprinted adult geese directed courtship behavior to ;oren- rather than other geese. @onrad ;oren-. ?obel ;aureate Animal Co!nition )ognition is the ability to thin$. 0hen applied in the context of animal behavior it refers to the ability of an animal to be aware of and ma$e judgements about its environment. Are animals conscious of themselves and their place in the world1 /o they feel pain, pleasure and sadness in the same way as humans1 Are the differences between animals and humans in cognitive ability absolute, as believed by /escartes, or a matter of degree1 !rofessor /onald Ariffin of !rinceton has been a proponent of the concept of cognitive ethology. This views conscious thin$ing as an inherent and essential part of the behavior of many non-human animals. )onscious thin$ing 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. /ifficult though they are to study the answers to 9uestions about animal cognition may profoundly affect our views on the treatment of animals and their welfare. 8ollow the lin$ to an article on animal thin$ing by !am ?oble entitled the Status of the Animal .ind Comm&ni ation )ommunication among animals relies upon their abilities to perceive sensory information. This may be visual, auditory or olfactory. /omestic animals perceive the world in a different fashion to us so may respond differently also. Vision - An obvious example of seeing the world differently relates to eye placement. %orses are set laterally, providing a wide range of monocular vision 2=35 degrees7 but a small :'-B' degree arc of binocular vision. n contrast, the cat has a much smaller arc of vision 2around 34' degree7 because of the placement of the eyes to the front but more than half of this is binocular vision. 0hen the relative visual acuity of domestic animals is compared, the ran$ing 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 fre9uencies than humans. )ats range of hearing is similar to humans. Olfactory senses - This is perhaps the most important sense of domestic animals in terms of communication. /ogs probably have the greatest olfactory ability of the domestic species. They are able to detect many compounds at 3C3'' the concentration of humans and for many wee$s after they were placed. #dors 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 2facial expressions, posture, grooming7 , and olfactory. These communication methods have varying importance between domestic species< Auditory - !igs have perhaps the most complex set of domestic animal vocal sounds - more than =' distinct types have been identified. %orses, 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 - n the horse, for example, the ear position can tell a lot about the animals disposition. *ars pointed bac$ 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 'lehmen res$onse. This is a curl of the lip when the urine of a female in estrus is smelled. n the cat, a high tail is a greeting or sign of being curious. >aised hac$les, and lips drawn bac$ in the dog denotes aggressionD if the ears are flat against the head and the tail is between the legs this signifies fear. !igs and horses show grooming behavior, as do many of the mon$eys and primates. Subordinate pigs groom dominant ones. %orses tend to groom horses of comparable ran$. )attle and cats spend long periods of time self-grooming but little time grooming others. Subordinate cattle will lic$ dominant cattle. Olfactory - #lfactory clues and scent serve to mar$ territory , show the way home, and distinguish individuals. Erine and feces are powerful means of olfactory communication. )ats and dogs also have anal sacs which are additional scent organs. #lfactory 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 li$e her own lamb. Animal "ehavior and the Animal Scientist.

/omestic animals display the same ranges of behaviors that would be seen in other animals and have both instinctive and learned behaviors. Enderstanding domestic animal behavior is not only fascinating and intellectually stimulating it is also crucial in their management, productivity and welfare. Behavior an# the #ete tion of animal #isease% #ften 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 recogni-e such as acute laminitis others can be 9uite subtle. An animal $ic$ing at its belly probably has an abdominal pain, while sweating in horses is a sign of acute pain e.g. colic. /ogs with ear infections often tilt their heads. *xcessive grooming or scratching is indicative of parasites or irritations. A change in the order in which an animal comes into the mil$ing parlor, particularly if a dominant cows hangs bac$, may indicate illness. Control of Bree#in! an# 'ee#in! 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 si-e of animals to reduce aggresion and so on. Trainin! n 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. 8or example the genetic predisposition of the border collie to herd is adapted by consistent training to increase s$ills 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 an# the Environment #bservation of animal behavior provides a great deal of information about the extent to which the animal copes with the environment in which they are $ept. t can also contribute to the design of better housing and management systems for farm animals. An obvious example is climate. Temperatures which are either too hot or too cold will lead to behavioral changes which are readily detectable. f dairy cows are standing in free stalls rather than feeding or lying down it probably indicates that the stalls are uncomfortable. f the environment of pigs is very barren and unstimulating this will lead to some aberrant behaviors such as aggresion and also to stereotypes. This is discussed further under the animal welfare section. Biblio!ra$h( )raig, Fames. 236437. /omestic Animal behavior< )auses and mplications for Animal )are and .anagement. !rentice-%allD, nc. *nglewood )liffes, ?ew Fersey %oupt, @atherine. 236637 /omestic Animal "ehavior for Geterinarians and Animal Scientists. owa State !ress, Ames, owa, The Traditions of Animal "ehavior< ?ature versus ?urture Etholo!( Although /arwin shifted the way we view animal behavior, the discipline also has a tradition that stretches before the time of /arwin 2/ric$amer and Gessey, 364:7. The field of ethology, which is the study of the evolution and functional significance of behavior, originated with ). #. 0hitman in the 34''+s. 0hitman coined the term instin t to describe the display patterns of pigeons. The etho!ram) a graph of the time course or switch points in a se9uence of behaviors, became a way of categori-ing species-typical behaviors. .any of these instincts are triggered by various environmental stimuli and von Eex$ull termed such triggers of instinctive stereotyped behaviors si!n stim&li. A classic stimulus triggers the courtship display of male three-spined stic$lebac$s fish. The enlarged belly of a female triggers the -ig--ag dance in male stic$lebac$ fish. The males use the dance to entice the female stic$lebac$ to enter the nest that the male has built. .uch of the wor$ of early ethologists was synthesi-ed by two ?obel ;aureates, ?i$o Tinbergen and @onrad ;oren-. ;oren- is noted for his wor$ on genetically programmed behaviors in young

and for studies on im$rintin!, during critical developmental periods in young. A classic example of imprinting occurs in young geese when they form an image of parent just after hatching. f the hatchlings first encounter a human such as ;oren-, they will imprint on him and follow him around as if he were their mother. A third ?obel ;aureate, @arl von 8risch, pioneered studies in bee communication and foraging. #ne of Tinbergen+s seminal contributions to "ehavior was to formulate a method studying animal behavior 2Tinbergen, 36:H7. This method forms the basis for how have structured material in this text. These issues are central to developing a philosophical approach to animal behavior. The ethological approach had a strong /arwinian tradition underlying its development. .uch of the wor$ in ethology was aimed at understanding the ultimate evolutionary reasons for behavior. Tinbergen listed four areas of in9uiry that could be used to understand issues of animal behavior. The following mnemonic can be used to remember these four areas A")/*8 I;ehrman, 36:5J< A ** Animal refers to the organisms. B ** Behavior refers to the observable actions of the organism. C ** Ca&sation refers to the proximate causes of behavior such as genes, hormones, and nerve impulses that control the expression of behaviors. D ** Develo$ment refers to the ontogeny of behaviors such as imprinting, or in the case of cognition, learning. E ** Evol&tion refers to the phylogenetic context in which behaviors are found. 8or example, the prevalence of parental care in birds, but not reptiles 2with some exceptions7 is an example of the taxonomic affiliations of some behaviors. ' ** '&n tion refers to the adaptive value or contribution that the behavior ma$es to fitness. +s( holo!( an# Behaviorism The ethological approach typified by the research of ;oren-, Tinbergen, and von 8risch 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 2e.g., ?orway rat7 in a controlled laboratory setting. )lassic wor$ by ". 8. S$inner lead to the development of the use of learning paradigms, and the S,inner Bo" remains an important tool in the field of animal psychology. 'i!&re -% A rat learns to press a bar in a S$inner "ox. 0ith each bar press the rat is rewarded with food. ;earning 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<

3. $er e$t&al $s( holo!( -- reception of environmental stimuli through the senses, and subjective perceptual interpretation of these sensory stimuli, =. $h(siolo!i al $s( holo!( -- an attempt to relate physiological properties within an organism to external behaviors 2e.g., measuring nerve impulse transmission in sensory and motor nerves7, H. f&n tionalism -- the study of the mind 2e.g., Fohn /ewey7 and how the mind operates. &. behaviorism -- the study of how accumulated experiences shape the behavior of the organism. The idea that an organisms is born a tabula rasa or 2blan$ slate7 upon which experiences accumulate and shape behavior is central to behaviorism. 5. animal $s( holo!( -- while initially related to the study of learning in model systems, the field of animal psychology in the present day encompasses a large body of wor$ related to cognition in a diverse group of animals. The Debate on Nat&re vers&s N&rt&re The field of *thology typified by the wor$ of Tinbergen, ;oren-, and von 8risch, and the broadly defined field of comparative psychology formed two drastically different schools of thought on the causes of behavior. 0e can compare and contrast their views to develop a deeper understanding behavioral analysis. The field of ethology, which originated in *urope, loo$ed to the genetic underpinnings of behavior. n contrast the field of comparative psychology, which originated in America, viewed behaviors as largely the product of the environment. /ifferences between the ethology and animal psychology led to a debate on the causes of behavior that has been captured in the often-9uoted phrase ,nature versus nature,. 0hat influences behavior -genes or environment1 The answer to this contentious debate cannot be put in terms of either genes or the environment, but must instead be loo$ed at in terms of a more complex interaction between genes and the environment. Behavioral E olo!( an# So iobiolo!( Students of Behavioral E olo!( have attempted to synthesi-e both the evolutionary traditions of *thology, and the mechanistic studies of )omparative !sychology. 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 loo$s at how organisms interact in their natural environments 2@rebs and /avies, 364B7. >esearchers are interested in both the mechanistic underpinnings of behavior, as well as the fitness conse9uences of behavioral traits. This tradition can be traced bac$ to Tinbergen and the four study areas 2)ausation, /evelopment, *volution and 8unction7. "ehavioral ecology is more broad than just a study of behavior, but also draws in issues of energetics and physiology 2e.g., )alow, 364B7. >ather than measure differences in survival and reproduction of behavioral traits, behavioral ecologists often use behavioral traits that maximi-e energy ac9uisition or foraging success as proxies for fitness traits. The development of optimal foraging during the B'+s and 4'+s has added a distinct theoretical perspective to the field of "ehavioral *cology. The newest approach to studying behavior involves a consideration of social systems in a diverse group of organisms. This field has ta$en off since the publication of So iobiolo!( by *. #. 0ilson 2364'7. "ecause some of these ideas have been applied to humans, the theory has been the target of much controversy. Sociobiology has a strong /arwinian tradition as it attempts to develop rules that explain the evolution of social systems and as such it. .ore recently, the field of Evol&tionar( +s( holo!( 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. %umans are considered subject to the same ,organic rules, that shape other organisms. ?eedless to say, this area is ripe for debate as researchers attempt to derive explanations for behaviors displayed by humans in modern society.

Eltimate versus !roximate )auses The dichotomy between *thology and )omparative !sychology with their concerns for adaptation and mechanism respectively, can be succinctly described as a concern for &ltimate versus $ro"imate a&ses. *rnst .ayr 236:37 describe the pursuit of those ultimate causes as a concern for the ,0hy Kuestions., 0hy does a bird give parental care1 0hy is a bee brightly colored1 n contrast, the pursuit of proximate causes is concerned with the way the world wor$s or the ,%ow Kuestions., %ow does a bat transmit echoes1 %ow do nerves carry impulses1 0here are memories stored1 Tinbergen+s four study areas also bloc$ out into ultimate versus proximate causation. 8or example, Tinbergen+s view of causation is concerned with !roximate )ausation, or mechanism. /evelopment is also considered to be in the category of proximate cause. %owever, evolution or phylogenetic context is s9uarely 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 2)urio 366&7. #ur study of animal behavior begins with a consideration of the ultimate causes of evolutionary change -- adaptation and natural selection. Ca&se) Develo$ment) Evol&tion) an# '&n tion Tinbergen+s brea$down can be used as a summary of the material covered thus far. prefer to ma$e the brea$down a little more detailed to include other approaches that have been added more recently by "ehavioral *cologists and Sociobiologists : .enes) E olo!() +h(siolo!() Develo$ment an# Learnin!) Evol&tion) an# So ialit(. This categori-ation 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. !aul Sherman 236447 would add yet another category to the list -- Co!nition. %owever, as cognitive theory is an outgrowth of development and learning, it will be included in those categories. "ehavioral *cology is undergoing a large-scale renaissance as researchers attempt to generali-e the classicallydeveloped ideas of !sychology and )ognitive !rocesses into wild populations 2>eal, 366&7. The first two subjects in the se9uence .enes and E olo!( will cover the basics of /arwinian natural and sexual selection as they apply to animal behavior. To cope with environmental variation, the organism evolves adaptations of $h(siolo!( that promote successful survival or reproduction. Such physiological changes could act at the level of endocrinology, neurophysiology, metabolism, or any of the myriad of proximate mechanisms that operate in an organism. These proximate mechanisms are used to help the organism cope with both abiotic 2e.g., the extremes of weather, navigation, etc.7 and biotic environmental factors 2e.g., the social environment, predation, etc.7. Additional components to an organism+s life are the #evelo$mental changes and learnin! that occur from ovumCsperm to maturity that are also adaptations to a particular way of life. 0hereas physiology operates in the very short term, development unfolds during the lifespan of an organism. 0ith an understanding of these genetic, ecological, physiological, developmental and cognitive processes in hand, we will be ready to tac$le the concepts of behavioral evolution. +h(lo!en( an# Constraints on the Evol&tion of Behavior Ep to this point, have operated under the premise that adaptation is the sole process that governs the evolution of behavior. %owever, in recent years, students of animal behavior have become more sensitive to the limitations of organic systems to change in an evolutionary sense. #rganisms may be well adapted, but limitations in organismal design constrain adaptation. n addition, organisms are also constrained by the effects of history or their own $h(lo!en(. /uring 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 ac9uisition of new adaptations 8unctional 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. )onsider a simple phylogenetic example ta$en from two lineages of vertebrates -- birds and mammals. All birds lay eggs, undoubtedly because the common ancestor of birds, some reptileli$e dinosaur, also laid eggs. %owever, most mammals bear live young because in the remote past a new $ind of mammal-li$e reptile evolved a different mode of life and passed this novel trait on to all subse9uent species in the linea!e or $h(lo!en(% A famous exception to this mammalian generali-ation includes the monotreme mammals of Australia, the platypus and echidna. t is thought that the monotremes branched off from the main stoc$ of mammals so early in the past that they retain the more ancestral mode of egg-laying reproduction. Such differences in reproductive mode 2egg-laying versus live-bearing7 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. *volution 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. n 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 mil$y substance which is secreted by part of their digestive system called the crop. "ecause both male and female birds have the crop, in theory both parents can evolve to produce a mil$y 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. n order to understand phylogenetic constraints that operate on other traits, we need a wor$ing $nowledge of the proximate mechanisms, as well as the process of natural selection. Accordingly, leave the discussion of such higher order macroevolutionary process for later chapters. So ietal an# C&lt&ral Evol&tion 8inally, leave the discussion of sociality until the very end, because it includes even more complex interactions that occur between organisms such as communication. The added complexity of sociality ma$es the study of behavior very rich indeed. A simple example will suffice. n developing our paradigm for animal behavior, 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 &lt&re introduces another mode of long-standing transmission of behavioral traits between generations. #ne need only wal$ into the nearest library to reali-e the impact of mass storage of human culture has on cultural transmission of culture. ;ibraries are a vehicle whereby information is passed on to subse9uent 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 &lt&ral transmission of information. Lour reading of this boo$ forms a $ind of cultural inheritance. .eneti Infl&en es in Animal Behavior )learly, genes significantly influence animal behavior. This is the only reasonable conclusion in cases where animals born and reared in isolation nevertheless develop age-appropriate, speciesspecific behaviors that they could not have possibly learned from other individuals. Such instincts that occur even in isolated animals include insect mating behaviorD courtship, nesting, and broodrearing behavior of pigeonsD the songs of some 2not all7 birdsD bird flightD and nut-crac$ing and nut-burying by s9uirrels. Animals are born ,$nowing, how to do certain things. *xperiments that rule out social learning and trial-and-error learning leave heredityMthat is, genesMas the only logical explanation for such behaviors.

!eople have long used artificial selection to produce animal breeds with desirable behavioral traits, such as dogs that herd sheep or hunt. Artificial*xperiments with honeybees have confirmed the relationship between genes and behaviors.selection can also shape the reactions of fruit flies 2/rosophila7 to light and gravity and the ability of rats to learn ma-es. Such results would not be possible if genes did not influence behavior. The fact that N rays and chemicals can induce mutations that alter behavior strengthens the lin$ to genetics. .utations have changed obstacleavoidance behavior of !aramecium and biological cloc$s and several behaviors in /rosophila. E"$erimental Evi#en e of .ene*Behavior Lin,s "reeding experiments confirm the relationship between genes and behaviors. 8or example, wor$er honeybees normally react to diseased or dead pupae by uncapping the honeycomb cell containing the pupa, dragging the pupa out, and removing it from the hive. This helps to prevent the spread of infections through the colony. *xperiments in which normal honeybees were crossed with bees that do not bring out their dead traced the behavior to two genes< one that induces wor$ers to uncap the diseased cell, and the other that induces the insects to remove the diseased pupa. %ybrids 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 /rosophila, breeding behaviors of cichlid fishes, food preferences in garter sna$es, bird migratory and nesting behaviors, and bird distress calls. +AVLOV) IVAN +ETROVICH /0123403-56 >ussian biologist who won the 36'& ?obel !ri-e in physiology for his demonstration of the idea of a ,conditioned reflex., !avlov trained dogs to drool at the sound of a bell by feeding them immediately after sounding the bell. LOREN7) 8ONRAD /039-403136 Austrian biologist who founded the study of animal behavior, or ethology. ;oren- said that animal behavior evolves in the same way as physical structures, such as wings. ;oren- shared the ?obel !ri-e in physiology with fellow ethologists @arl von 8risch and ?i$olaas Tinbergen. 8inally, 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. Aenes encode proteins, nothing moreD but through proteins, they can influence behavior. Aggression and sexual behavior, for example, are influenced by testosterone, and testosterone is synthesi-ed by en-ymes, which are proteins encoded by deoxyribonucleic acid 2/?A7. Thus one can see how genes would influence these behaviors. All behavior, furthermore, depends on chemical signals 2neurotransmitters7 that are released by one neuron and bind to receptors on the next neuron. ?eurotransmitters, too, are synthesi-ed by en-ymes encoded by /?A, and their receptors are proteins as well. ?eurotransmitter levels control mood and probably aspects of personality. The list goes on and on. ndeed, it is impossible to see how genes could not play a role in behavior.