Important words and concepts from Chapter 51, Campbell & Reece, 2002 (3/25/2005):

by Stephen T. Abedon ( for Biology 113 at the Ohio State University



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(1) Chapter title: Behavioral Biology

(a)                    [glossary of terms used in animal behavior and evolution (Behavioral EcologyJane BrockmannUniversity of Florida)]

(b)                    [behavioral biology (Google Search)] [index]




(2) Behavioral ecology

(a)                    We expect natural selection to attempt to maximize the Darwinian fitness associated with the traits possessed by organisms

(b)                    The goal of a behavioral ecologist is to explain behaviors in terms of their impact on Darwinian fitness

(c)                    As with any trait (given an “adaptationist paradigm”), the behavior of an organism, too, we expect to be Darwinian-fitness enhancing

(d)                    That is, behaviors that reduce Darwinian fitness we predict will be lost from gene pools while behaviors that increase Darwinian fitness we expect to increase in frequency within populations

(e)                    Note, however, that while behaviors often have underlying genetic components, the ability of organisms to learn complicates our understanding of behavioral evolution; nevertheless, we have an expectation that organisms will have a tendency to behave in a manner that enhances their Darwinian fitness

(f)                      [behavioral ecology (Google Search)] [nebraska behavioral biology group] [animal behaviour--zoological record (Biosis)] [index]

(3) Behavior

(a)                    What is behavior?

(i)                      Defined extremely broadly, behavior is how organisms act in response to environmental stimuli, with the word "act" (as well as the phrase "environmental stimuli") left somewhat ambiguous

(ii)                    Clearly organisms act or react physiologically to all sorts of environmental stimuli (e.g., Escherichia coli's modification of gene expression in response to low levels of the sugar glucose and high levels of sugar lactose)

(b)                    The study of behavior becomes interesting when behaviors

(i)                      are not obviously in the organism's best interest

(ii)                    when the cost of the behavior is high even given that ultimately the behavior serves the organism's Darwinian interests (e.g., in terms of energy required to learn or display the behavior, or when the maintenance of costly anatomical features, e.g., a large brain, is necessary for the display of the behavior), or

(iii)                   when the behavior appears to be overly simplistic but still sufficient to get the job done (e.g., FAPs)

(c)                    We would say that an organism's behavior is optimized when the organism's Darwinian fitness is optimized given the display of that behavior relative to the display of some alternative behavior, including the display of no behavior at all (“If you choose not to decide, you still have made a choice.”)

(d)                    Behavior typically influences the acquisition of energy, nutrients, sex, help in child rearing, removal of ectoparasites, survival, establishment and maintenance of dominance hierarchies, etc.

(e)                    Note that whereas behavior is traditionally studied with animals, unless defined in some way that excludes all other kingdoms (e.g., requiring muscles or a nervous system) then behavior is the province of members of all kingdoms (plus their viruses); nevertheless, behavior obviously is displayed with a greater complexity by animals so the animal-emphasis of behavioral ecology is quite justified

(f)                      [behavior (Google Search)] [index]




(4) Ultimate causation

(a)                    When we question why a behavior occurs (or, for that matter, why any trait exists), we are posing a question of ultimate causation

(b)                    Ultimate causation is simply another way of saying optimization of Darwinian fitness

(c)                    The ultimate cause of a behavior is its increase in frequency within a population due to the greater Darwinian fitness that results from organisms engaging in that behavior

(d)                    Thus, when questioning ultimate causation, one is simply asking why a behavior evolved

(e)                    The hallmark of the discipline called behavioral ecology is its emphasis on determining the ultimate causation of behaviors

(i)                      Understanding the ultimate causation of some behaviors can be very straightforward (a female songbird mates to have babies)

(ii)                    Interesting questions in behavioral ecology are those that are less straightforward (e.g., a female songbird mates with someone other than her lifetime mate or why a bird would push her own eggs out of her own nest)

(f)                      [ultimate causation (Google Search)] [index]

(5) Proximate causation

(a)                    In contrast with ultimate causation, proximate causation involves the mechanics of how a behavior occurs including

(i)                      how stimuli are received

(ii)                    how received stimuli are translated into a response, and

(iii)                   the mechanisms underlying the response (i.e., the behavior)

(b)                    A behavioral ecologist assumes that the proximate cause of a behavior is simply the mechanism underlying the means by which a behavior is manifest

(c)                    "Thus, the 'how' and 'why' questions about animal behavior are related in their evolutionary basis: Proximate mechanisms produce behaviors that ultimately evolved because they increase fitness in some way."

(d)                    As such, a behavioral ecologist is interested in proximate causation only to the extent that these mechanisms serve to constrain the evolution of specific behaviors

(e)                    It should not be overlooked that understanding how behaviors are constrained by an organism's anatomy and physiology is highly relevant; however, otherwise becoming mired down in the details of the anatomy and physiology of a behavior can be somewhat distracting to one's understanding of the ecology of a behavior, so proximate causation is frequently accepted as a given (i.e., the behavior happens so the organism must be anatomically, physiologically, and mentally capable of expressing the behavior)

(f)                      [proximate causation (Google Search)] [links related to animal perception (Animal Learning & Cognition)] [index]

(6) Umvelt (supplemental discussion)

(a)                    This is the term used to describe the means by which proximate causation underlies the display of a given behavior, that is, the anatomical, physiological, and environmental (?) context underlying and allowing a behavior to occur

(b)                    [umvelt (Google Search)] [how animals see (Animal Learning & Cognition)] [index]

(7) Ethology

(a)                    Behavioral ecology as a discipline had its roots in the more proximate causation-concerned field of ethology

(b)                    Ethology, however, is less concerned about learning and more concerned about innate, not-learned behaviors

(c)                    "One of the major findings of ethology was that animals can carry out many behaviors without ever having seen them performed. In other words, many behaviors are innately programmed. And while such behaviors seem purposeful because they are clearly beneficial, they are carried out in ways that show the animals are unaware of the significance of their actions."

(d)                    These are examples of innate behaviors:

(e)                    [ethology (Google Search)] [ethology and evolution on the web (The International Society for Human Ethology)] [index]

(8) Fixed action patterns (FAP) [sign stimulus, releaser]

(a)                    A fixed-action pattern is a series of behaviors that are both innate and, typically, are completed in full once initiated (even if their completion is laughably inappropriate)

(b)                    The initiation of a fixed action pattern is in response to an external sensory stimulus known as a sign stimulus or a(n innate) releaser

(c)                    The reason why FAPs are not always appropriate is because the releaser for the action is often a proximate correlate to the stimulus ("a limited subset of the available sensory information") that while the FAP is ultimately (i.e., evolutionarily) a response to some full stimulus

(d)                    "We can think of a FAP as the innate ability of an animal to detect a certain stimulus associated with the animal's umvelt, combined with an innate behavioral program that is activated by the stimulus to direct some kind of motor activity."

(e)                    "In the case of a FAP, the animals are behaving more like robots."

(i)                      However, the important thing to keep in mind is that the FAP under natural conditions typically represents the energetically minimal investment in the display of a behavior that otherwise is good enough (i.e., evolution is not just optimizing the outcome of the behavior but also optimizing the efficiency of the behavior such that behavioral outcome for energy invested is maximized)

(ii)                    "The ability to confront novel stimuli, learn about them, and adjust behavior is a hallmark of both intelligence and self-awareness. The evolution of intelligence is costly, in both the development of the neural tissue necessary to process the information and its metabolic maintenance. In addition, the evolution of intelligence requires dramatic changes in life history patterns, such as long juvenile phases and high parental investment per offspring. For most species, these costs, measured as reductions in reproductive fitness, far outweigh the costs of an occasional inappropriate use of FAPs, and extensive intelligence has not evolved in many animal groups."

(iii)                   "Some ethologists have suggested that the use of simple cues to release programmed behavior prevents an animal from wasting time processing or integrating a wide variety of input. Perhaps a better way of interpreting the situation has to do with the limitations of innate behavior and how it evolved. A frog's sensory-neural network for detecting movement is probably much less complex than the apparatus that would be necessary to rapidly distinguish a fly from another object of similar size. In any case, simple cues usually work quite well in an animal's normal sensory world, though not in the experimental world ethologists often create."

(f)                      [egg rolling animated gif (1.8 MB file), chick hatching movie (600 KB) (Behavioral Control Systems)] [fixed action patterns, sign stimulus, innate releaser (Google Search)] [stimuli and innate behavior (Bio 170 – Animal Behavior – Barry Sinervo)] [index]




(9) Learning

(a)                    Learning is synonymous with behavioral modification such that, ideally, a behavior is further optimized by the change

(b)                    Learning can optimize behavior most obviously in a short term sense; however, for behavior to be truly optimized, it must be optimized in a Darwinian sense

(c)                    "Nearly all biologists today agree that most behavior is a consequence of genetic and environmental influences. Even though an animal may not have to witness a FAP because the basic behavior is innate, learning is still involved. Most FAPs improve with performance, as animals learn to carry them out more efficiently."

(d)                    Learning may be differentiated into a number of different types and associated concepts:

(i)                      Maturation (not learning at all)

(ii)                    Habituation (learned ignoring)

(iii)                   Imprinting (learned sign stimulus)

(iv)                  Associative learning

·        Classical conditioning (Pavlov's dog)

·        Operant conditioning (trial and error learning)

(v)                    Observational learning (copying)

(vi)                  Play (practicing)

(vii)                 Insight learning (assessment of novel situations)

(e)                    [animal learning (Google Search)] [index]

(10) Maturation

(a)                    Learning must be operationally distinguished from maturation

(b)                    That is, just because an animal improves its performance of a behavior with time does not mean that the animal is improving through learning; it could instead mean that the animal was physiologically or anatomically unable to properly or fully display the behavior until they have reached an appropriate developmental stage

(c)                    Such development-associated improvements in behaviors occur via maturation

(d)                    [animal maturation and learning (Google Search)] [index]

(11) Habituation

(a)                    Habituation is one means by which animals adapt behaviorally to their environment (i.e., learn)

(b)                    Habituation is the learned ignoring of a stimulus

(c)                    This occurs when an animal learns that a stimulus is not appropriately correlated with the expected full stimulus

(d)                    Thus, habituation is a means of, for example, energy conservation employed when proximal stimuli are not followed by subsequently expected stimuli

(e)                    Habituation in seals:

(f)                      [animal habituation and learning (Google Search)] [index]

(12) Imprinting (critical period)

(a)                    Imprinting is a FAP to a stimulus that is learned rather than a stimulus that is innately known

(b)                    Two things distinguish imprinting from other types of learning

(i)                      An imprint, once learned, is never forgotten (i.e., imprinting is irreversible)

(ii)                    An imprint can only be learned during a critical period, which is defined (somewhat tautologically) as the limited time during which an imprint may be learned

(c)                    "While a critical period and irreversibility characterize imprinting, it now recognized that these phenomena are not always rigidly fixed." That is, it is possible, though not easy, in some situations/species for an animal to eventually modify its behavior to disregard imprinting

(d)                    See Figure 51.9, Imprinting

(e)                    [imprinting, definition] [imprinting, imprinting "critical period" (Google Search)] [index]

(13) Associative learning

(a)                    Associative learning is the association of one stimulus with another

(b)                    See classical conditioning and operant conditioning as examples

(c)                    [associative learning (Google Search)] [index]

(14) Classical conditioning

(a)                    Classical conditioning is the occurrence of associative learning following repeated, correlated exposure to two stimuli

(b)                    In classical conditioning each stimulus becomes equivalent in terms of the behavioral response

(c)                    Thus, otherwise irrelevant correlates to relevant stimuli can come to induce identical responses (e.g., the ringing of bells with the smell/taste of food; the opening of refrigerators or the rustling of plastic bags and the whistling response by guinea pigs used to being fed “fresh” produce)

(d)                    [classical conditioning (Google Search)] [index]

(15) Operant conditioning

(a)                    Operant conditioning is trial-and-error learning

(b)                    Operant conditioning works (in an ultimate, Darwinian sense) so long as an animal possesses a reasonable means of distinguishing good from bad

(i)                      Using these terms in their ultimate, Darwinian sense, e.g., a good thing results in more offspring while a bad thing results in less offspring

(c)                    This means of distinguishing good from bad requires either a reasonable idea of the future consequences of one's actions (e.g., as humans supposedly possess) or some kind of physiological indicator of good and bad, which in animals is represented (probably) as pleasure and pain

(d)                    Thus, animals continue to do things that give them pleasure while refraining from doing things that cause them pain

(e)                    Keep in mind that the reaction of pleasure or pain is itself a product of natural selection, such that things that tend to enhance Darwinian fitness tend to be pleasure-filled while those things that tend to diminish Darwinian fitness are painful

(i)                      Of course, if an environment changes then the correlation between painful and diminishment of Darwinian fitness or pleasurable and enhancing of Darwinian fitness may be lost

(f)                      Note that while classical conditioning is the association of two stimuli such that an animal anticipates a result (one the stimuli; e.g., food) given the stimulation by the other (e.g., a bell), in operant conditioning the second stimulus in a sense is internal rather than external: the animal associates stimuli either with pleasure or with pain and learns to seek out those stimuli that elicit the former and avoid those that elicit the latter; thus, classical conditioning involves the association of two external stimuli while operant conditioning involves a learned association between an external stimulus and an internal state

(g)                    See Figure 51.11, Operant conditioning

(h)                    [operant conditioning (Google Search)] [index]

(16) Observational learning

(a)                    Observational learning is the copying of the actions of others

(b)                    The assumption is that others, by having survived, etc., possess behaviors that are worth knowing

(c)                    Note that by this logic the worth of another's behavior (i.e., whether that behavior is worth copying) diminishes with the age of the other such that while the behavior of an older individual may be worth copying

(i)                      The behavior of a same-aged individual is less worthwhile

(ii)                    The behavior of a younger individual has the least worth of all

(d)                    [observational learning (Google Search)] [index]

(17) Play

(a)                    Play is essentially practice

(b)                    The more we practice, the better we get at something (which ideally enhances our survival or reproductive success); this "getting better" can be in terms of the acquisition of skill or in the development and maintenance of musculature that will be relevant to possess once even extended parental care is no longer available

(c)                    Consequently, the finding of pleasure in play is our body's way of making sure that we practice

(d)                    Note that play is an expensive way to learn:

(i)                      It requires a big brain that is capable of learning complex behaviors

(ii)                    To the extent that play requires energy while exposing individuals to risk, play is a costly behavior in and of itself

(e)                    [play (with over 30 million hits, I suspect that “play” is not the most useful search strategy) (Google Search)] [index]

(18) Insight learning (supplemental discussion)

(a)                    This is the performance of a novel behavior that occurs as a consequence of thinking through the behavior and at least some of its consequences (particularly short-term consequences) before initiating the behavior, i.e., basically figuring something out for oneself without continuous, direct environmental feedback

(b)                    Only the very smartest animals are capable of insight learning

(c)                    [insight learning (Google Search)] [index]




(19) Moving and timing behaviors

(a)                    A number of animal behaviors might (for convenience) be described as movement and timing behaviors (a category I've made up essentially to serve as a heading for the discussion that follows); these include

(i)                      Rhythmic behaviors (supplemental discussion)

(ii)                    Kinesis

(iii)                   Taxis

(iv)                  Migration

(b)                    [(Google Search)] [index]

(20) Rhythmic behaviors (supplemental discussion)

(a)                    Animals typically display particular behaviors at different times of the day or times of the year

(b)                    The mechanisms underlying these rhythmic behaviors may be categorized as endogenous or exogenous (coming from within the animal and coming from the animal's environment, respectively)

(c)                    Typically the exogenous signal is light (i.e., the stuff that comes from the sun)

(d)                    For example, a typical animal will be able to almost-display rhythmic behaviors at the appropriate times without exogenous signals (e.g., if deprived or light or changes in light) but will only not be able to display behaviors at appropriate times with high accuracy if deprived of the exogenous signals

(e)                    Note that this endogenous-exogenous system allows animals to innately display appropriate behaviors at nearly appropriate times independently of fallible exogenous signals, but still adapt to changes in exogenous signals, e.g., as day lengths change with the seasons

(f)                      [rhythmic behaviors, circadian rhythm, chronobiology (Google Search)] [index]

(21) Kinesis

(a)                    Kinesis is a movement behavior such that movement randomly occurs given the presence of a stimulus; in practice, an organism will tend to settle down in a region that is preferred by tending to move particularly when not present in the preferred location

(b)                    Kinesis is a mechanism of movement that involves activity only so long as a stimulus is present (or, alternatively, only when a signal is absent)

(c)                    For example, an animal might move only when the environment is in some way unfavorable; this movement does not occur in a specific direction but instead is randomly directed, with attainment and maintenance of the animal within a more favorable environment occurring simply because less movement occurs when the movement-associated stimulus is not present

(d)                    [Kinesis is the animal equivalent of the answer to “Why do so many leaves collect in my pool?” The answer is that fallen leaves often may be blown by the wind. However once they are sufficiently wet they no longer may be blown around. These leaves are capable of more or less freely moving about until they land in your pool. Thus, they tend to collect in bodies of water.]

(e)                    [A kinesis-like mechanism probably also explains what radio station you listen to. That is, as you are flicking through the “dial,” your tendency is to change stations when a station plays a song (or whatever) that you do not like. Therefore it is the stations that most frequently play things that you don’t (or least) want to listen to that will be changed the most. Thus, chances are you the station you listen to is more the one that plays the music you find least objectionable rather than the one that plays the music you enjoy the most.]

(f)                      [One likely can generalize the above scenario to all sorts of aspects of your life from produce to mate choice. For instance, the above arguments probably also help to explain the American diet: We are far more interested in obtaining foods that we don’t find objectionable than obtaining foods that we find absolutely wonderful. As a consequence, it is very difficult to find absolutely wonderful food in America.]

(g)                    [animal kinesis (Google Search)] [index]

(22) Taxis

(a)                    Taxis, by contrast with kinesis, is directed movement either toward or away from a stimulus

(b)                    [animal taxis (Google Search)] [index]

(23) Migration

(a)                    Migration is the regular (annual) movement of animals over long distances

(b)                    Migrating animals find their way via one of three mechanisms

(i)                      Piloting ("map")

(ii)                    Orientation ("compass")

(iii)                   Navigation ("map" and "compass")

(c)                    [animal migration (Google Search)] [index]

(24) Piloting

(a)                    Piloting is the movement from one landmark to another such that terrain remains familiar throughout the migration

(b)                    Because of the requirement that an animal have essentially memorized the directions for the entire trip (an internal map), piloting is typically employed only for relatively short trips (or trips made over well land-marked land, e.g., over a well-worn path)

(c)                    [(Google Search)] [index]

(25) Orientation

(a)                    Orientation involves straight-line travel in a direction that is globally oriented (e.g., flying by compass)

(b)                    Compasses are not necessarily artifacts (i.e., man-made devices) but may also be small, sensory magnets found within an animal’s head

(c)                    Alternatively, many animals appear to orient using the position of the sun (or stars) and innate knowledge of the time of the day

(d)                    [animal orientation Google Search)] [index]

(26) Navigation

(a)                    Navigation involves possessing both a map and a compass

(b)                    Maps are not necessarily paper things, i.e., one can possess a “map” within one's head

(c)                    [animal navigation Google Search)] [index]




(27) Foraging (generalists, specialists)

(a)                    Foraging is the behavior(s) an animal employs to feed

(b)                    "Food habits are a fundamental part of an animal's niche and may be shaped in part by competition with other species."

(c)                    Animals may be categorized in terms of their foraging behaviors into generalists and specialists

(d)                    Specialist tend to be better at acquiring specific kinds of food, employing morphological adaptations as well as behavioral adaptations and culture to maximize their yield of specific foods

(e)                    Generalists are not as good at acquiring any particular kind of food, but generalists potentially have available to them many more kinds of food; "Generalists cannot be as efficient at securing any one type of food, but they have the advantage of having other options if a preferred food becomes unavailable."

(f)                      [foraging, foraging behaviors (Google Search)] [index]

(28) Search image

(a)                    Generalists, too, can specialize in the acquisition of specific food stuffs, particularly by temporarily specializing

(b)                    When specializing the generalist employs a specific search image which is a behavioral algorithm for catching/obtaining a certain kind of food

(c)                    This search image will be employed, and even improved upon, until a decline in the abundance of that food stuff leads to literally diminishing returns; then, rather than suffering without adequate food supplies, the generalist can simply switch to a new food type and new search image

(d)                    [animal "search image" -engine (Google Search)] [index]

(29) Optimal foraging

(a)                    What makes one food type and search image preferable to another?

(b)                    Basically, ideally, an organism chooses to consume the food that is easiest to acquire on a per-unit-nutrient basis and in optimal foraging the benefits of a given forage essentially are optimized relative to costs

(c)                    Thus, for example, slow, tasty, and abundant things are consumed preferentially to fast, rare, obnoxious things

(d)                    Organisms that perform such a cost-benefit analysis are said to be optimizing their foraging

(e)                    "Optimal foraging theory predicts that natural selection will favor animals that choose foraging strategies that maximize the differential between benefits and costs. Benefits are usually considered in terms of energy (calories) gained. However, other optimization criteria, such as specific nutrients, are sometimes more important than energy. Costs or tradeoffs associated with foraging consist of the energy needed to locate, catch, and eat food; the risk of being caught by a predator during feeding; and time taken away from other vital activities, such as searching for a mate."

(f)                      As a consequence of the complexity of the cost-benefit analysis of optimal foraging behavior, an organism does not necessarily do all necessary calculations in its head prior to subduing food, but instead has certain rule-of-thumb behavioral tendencies representing evolutionary algorithms defining what to eat under what circumstances; it is typically these rule-of-thumb behavioral tendencies that are optimized in optimal foraging behavior

(g)                    ["Behavioral ecologists use (many) factors to predict how an animal will forage given a particular set of conditions. Their goal is not to test whether animals in fact forage optimally, but to use the expectations of optimality as a guide to organizing research and generating testable predictions. When their predictions are borne out, researchers come closer to understanding the major factors that shape an animal's foraging behavior. When their predictions are not borne out, they have still made progress because they know they must consider additional factors. Predictions in optimal foraging studies are usually quantitative; they are frequently based on direct measurements of the calories an animal must expend to secure particular food items and the calories it gains by doing so. Numerous studies of many species show that animals tend to modify their behavior in a way that keeps their overall ratio of energy intake to energy expenditure high. Their ability to do this is sometimes quite surprising. The smallmouth bass is somehow able to factor in all the relevant variables and forage in a highly efficient manner, switching between minnows and crayfish as conditions change. The proximal mechanisms responsible for this process are not known. They must include innate cues, but experience undoubtedly modulates behavior."]

(h)                    [optimal foraging (Google Search)] [optimal foraging theory (Bio 170 – Animal Behavior – Barry Sinervo)] [index]




(30) Social behavior

(a)                    Social behavior is interactions between two or more organisms, typically animals, usually conspecifics

(b)                    Some animals are more social than others, with the minimal social behavior necessary between sexually reproducing animals being that associated with mating

(c)                    "The complexity of behavior increases dramatically when interactions among individuals are considered. Aggression, courtship, cooperation, and even deception are part of the behavioral landscape of social behavior. Social behavior has both costs and benefits to members of those species that interact extensively."

(d)                    [social behavior (Google Search)] [index]

(31) Sociobiology

(a)                    Sociobiology is the application of evolutionary theory to the study of social behavior (just as behavioral ecology is the application of evolutionary theory to the study of animal behavior)

(b)                    “Social scientists regard culture – everything from eating habits to language – as an entirely human invention, one that develops arbitrarily… Sociobiologists, by contrast, emphasize that learned behavior, and indeed all culture, is the result of psychological adaptations that have evolved over long periods of time. Those adaptations, like all traits of individual human beings, have both genetic and environmental components… Social science… promotes erroneous solutions, because it fails to recognize that Darwinian selection has shaped not only human bodies but human psychology, learning patterns and behavior as well.” (Randy Thornbill and Craig T. Palmer, 2000, Why men rape. The Sciences 40(1 – January/February): 30-36) (The Sciences is a publication of the New York Academy of Sciences)

(c)                    [sociobiology (Google Search)] [Behavioral Ecology and Sociobiology (a journal) (Springer)] [index]




(32) Agonistic behavior

(a)                    Aggressive behavior between conspecifics usually involves fighting over a limiting resource such as food, water, space, or mates

(b)                    Depending on the importance of the resource as well as its scarcity, agonistic behavior can range from all-out fighting to the death to much safer ritualistic behavior

(c)                    Animals typically lack action-at-a-distance weapons such as those possessed by humans (e.g., guns) so risk injury or even death whenever they engage in aggressive behavior

(d)                    Consequently, animals often avoid fighting unless there is a sure indication that they will win without incurring injury and if the resource is worth fighting over

(e)                    Animals often possess sophisticated rituals in which they attempt to bluff their opponent into backing down, and animals also often have a good sense of when to retreat as losers from an otherwise hopeless, potentially injurious cause

(f)                      Agonistic displays by a human female and a mandrill male:


(h)                    [agonistic behavior (Google Search)] [index]

(33) Dominance hierarchies

(a)                    One way in which agonistic behavior is avoided is by knowing one's place in the scheme of things and then avoiding any behaviors that might be interpreted as threatening to those possessing a higher rank

(b)                    Animals that are higher on dominance hierarchies risk injury to attain that status (and often must commit large amounts of energy to maintaining that status), but benefit by gaining preferred access to food or mates for themselves and also potentially for their offspring

(c)                    Thus, a dominance hierarchy represents an institutionalized (though not necessarily completely static) snapshot of the results of previous agonistic behavior, and are advantageously maintained by everyone to the extent that previous fighting need not be repeated

(d)                    [dominance hierarchies (Google Search)] [index]

(34) Territoriality (home range)

(a)                    Another way that all-out fighting is avoided is by institutionalizing conflicts into territories (dominance hierarchies might also be considered to be territories, ones where social position is defended rather than space)

(b)                    A territorial individual will typically defend a specific area particularly against intrusion by conspecifics

(i)                      Conspecifics are individuals that an individual is most directly in competition with, against which an individual possesses a reasonable chance of winning a fight, and with which one's mate might fool around

(ii)                    Territories that are not defended are instead called home ranges

(c)                    Territories may be permanent or temporary, may be defended by one individual or many, and may be established to guard space, food, mates, etc.

(d)                    Territory owners are more likely to win fights against would-be trespassers; this is probably because the owner

(i)                      Has more to lose and therefore is willing to risk more to win a fight (since the owner but not the trespasser has learned the territory)

(ii)                    May be an older, more experienced fighter (since younger, less experienced fighters presumably are less likely to have acquired a territory)

(e)                    In many species territories are indicated by more than just agonistic behavior, including expressions of sound and scent intended to warn off potential trespassers

(f)                      Note that territoriality represents an uneven partitioning of resources that can have the effect of minimizing the excursion of populations beyond an ecosystem's carrying capacity for that organism

(i)                      (thus, one would predict that K selection and territoriality would often go hand in hand; we will consider K selection in some detail in the next chapter)

(g)                    [territoriality (Google Search)] [index]




(35) Mating behavior

(a)                    "All aspects of reproductive behavior receive extensive attention from behavioral ecologists. The reason is that researchers can often determine the number of young an individual produces, which comes very close to determining an animal's fitness. The correlation between measurable behaviors and fitness may not be as strong for other aspects of behavioral ecology, such as optimal foraging."

(b)                    [mating behavior (Google Search)] [index]

(36) Courtship

(a)                    "In many animals, potential partners must go through a complex courtship interaction, unique to the species, before mating. This complex behavior often consists of a series of fixed action patterns, each triggered by some action of the other partner and initiating, in turn, the other partner's next required behavior. This sequence of events assures each animal not only that the other is not a threat, but also that the other animal's species, sex, and physiological condition [e.g., receptivity to mating] are all correct."

(b)                    Part of courtship behavior can additionally involve some degree of assessment by one individual of the other individual

(c)                    In addition to assuring that matings are successful in the sense that they produce viable offspring, an animal additionally is motivated to increase the likelihood of the success of offspring by finding, via courtship, mates that have the best genes or behaviors, i.e., animals, given a choice, like to "marry up"

(d)                    [courtship behavior (Google Search)] [The Mating Habits of Earthbound Humans (a mockumentory)] [index]

(37) Parental investment

(a)                    An animal that is destined to invest a lot of time and energy in the parenting of offspring would prefer to mate with individuals who bring more to the mating rather than less

(b)                    Typically, especially given internal fertilization, the gender that invests the most in offspring is the female

(c)                    This investment can come in the form of large gametes, internal development of the young, or raising the young following birth or hatching

(i)                      Most females contribute to a least one of these parental investments

(ii)                    Many males do not (though, to be fair, there are many males out there that contribute to the sheltering and nourishment of females, the sheltering or internal development of young, and the raising of young)

(d)                    Sexual selection is the means by which a gender (typically males) either compete directly over a female or compete for a female's favorable attention; these battles in turn select for secondary sexual characteristics that aid in the fighting of battles over females or the attracting of females

(e)                    In some species, e.g., humans, both parents often invest mightily in the raising of offspring and sexual selection consequently impacts directly on both genders

(f)                      [parental investment (Google Search)] [index]

(38) Mating systems [promiscuous, monogamous, polygamous, polygyny, polyandry]

(a)                    An animal's mating system refers to how animals pair up for mating

(b)                    Promiscuous mating systems exist particularly where one parent (e.g., the guy) does not participate at all in the raising of offspring whereas monogamous relationships occur particularly when two parents share the raising of children

(c)                    Various mating-system types include:

(i)                      Promiscuous = low likelihood of subsequent mating with same individual

(ii)                    Monogamous = high likelihood of subsequent mating with one individual

(iii)                   Polygamous = high likelihood of subsequent mating with more than one individual

(iv)                  Polygyny = one male mates with several females

(v)                    Polyandry = one female mates with several males

(d)                    It is important to keep in mind that a mating system usually exists to serve some ultimate cause, i.e., enhancement of both male and female fitness

(e)                    [mating systems, promiscuous mating systems, monogamous mating systems, polygamous mating systems, polygyny, polyandry, (Google Search)] [index]




(39) Altruism

(a)                    Altruism is cooperative behavior in which the actor's Darwinian fitness is reduced by the behavior

(b)                    Particularly, this is when the actor's individual (as opposed to inclusive) fitness is reduced by the behavior

(c)                    [altruism (Google Search)] [index]

(40) Cooperative behavior

(a)                    Cooperative behaviors are acts that increase the Darwinian fitness of others

(b)                    These same acts may or may not increase the Darwinian fitness of the actor, or may even decrease the fitness of the actor

(c)                    Since natural selection favors individuals who increase their own fitness (i.e., not necessarily that of other's), it is not always obvious nor easy to understand either why cooperative behavior occurs or how it might have evolved

(d)                    The evolution of cooperative behavior is typically considered to result as a consequence of either

(i)                      Kin selection

(ii)                    Reciprocal altruism

(iii)                   Group selection

(iv)                  Avoidance of punishment

(e)                    ["cooperative behavior" ecology (Google Search)] [index]

(41) Kin selection

(a)                    Kin selection is another way of saying that it pays to help those to whom we are most closely related

(b)                    Why?

(i)                      Individuals with which an individual is closely related share genes including genes that code for cooperative behavior

(ii)                    Increasing the Darwinian fitness of relatives therefore can serve to increase the Darwinian fitness associated with cooperative behavior

(c)                    Note that key to kin selection is the ability to distinguish non-relatives from relatives, either by recognition of some kind or via correlates such as when relatively non-mobile individuals tend to be more closely related to individuals who are near to them than they are to individuals who are a greater distance from them

(d)                    Kin selection essentially is cooperative behavior among kin (relatives) resulting in greater Darwinian fitness for these relatives in comparison to groups that do not mutually cooperate; the basis of kin selection is that relatives share genes and therefore that enhancing the fitness of a relative to some degree is equivalent to enhancing ones own Darwinian fitness (and, in particular, it enhances the inclusive fitness associated with ones genes/alleles)

(e)                    The most obvious form of kin selection is the caring of offspring by parents, i.e., it pays for parents to care for their offspring because their offspring share genes with them (that is, any allele that coded for not caring for otherwise helpless offspring would quickly go extinct)

(f)                      [kin selection (Google Search)] [index]

(42) Inclusive fitness

(a)                    Inclusive fitness is the idea that the fitness associated with an individual's genes is a function not just of the Darwinian fitness of the individual, but additionally the Darwinian fitness associated with the same alleles found in relatives

(b)                    Thus, as far as your genes are concerned, your survival (and consequent reproduction) is no more important than the survival of two full siblings (i.e., brother or sister; with each you share half of your genes) or eight first cousins (with each you share 1/8 of your genes); note that we are implicitly assuming that all considered individuals have identical reproductive potentials, e.g., are of the same age and health, etc.

(c)                    [inclusive fitness (Google Search)] [index]

(43) Coefficient of relatedness

(a)                    The coefficient of relatedness is the means by which these fractional-sharing-of-genes ideas are formalized

(b)                    The coefficient of relatedness is the proportion of alleles in two individuals that are identical by decent (the calculation of which can get messy if inbreeding has occurred)

(i)                      For example, half of your genes came from your mom and, on average, each of your siblings shares half of these genes (since the same is true for your father, one-half * one-half + one-half * one-half = one-half, i.e., you share half of your genes with each full sibling)

(c)                    [coefficient of relatedness (Google Search)] [index]

(44) Reciprocal altruism

(a)                    Reciprocal altruism is a means by which cooperative behavior can evolve even between non-relatives

(b)                    In reciprocal altruism, one individual performs a cooperative act in the (evolutionary) hope that individual who is the recipient of the act will cooperate in return

(c)                    Reciprocal altruism likely requires a means by which individuals can distinguish reciprocators from those who refuse to cooperate in return

(d)                    [reciprocal altruism, prisoner's dilemma (Google Search)] [prison project home page (site dedicated to the iterated prisoner’s dilemma, a metaphor for the evolution of cooperation via, particularly, reciprocal altruism) (LifL)] [index]

(45) Group selection

(a)                    Group selection is a means by which cooperative behavior might evolve, though chances are that in the real world it makes little contribution except, perhaps, in the evolution of culturally learned behavior that is cooperative

(b)                    The basic idea is that two groups are competing either over resources or directly fighting against each other; if in one group individuals cooperate whereas in the other they do not, then the overall fitness of the cooperating group will exceed that of the non-cooperating group, thus driving the latter, along with its non-cooperative behavior, to extinction

(c)                    For example, humans even within large groups are very willing to put aside differences to fight against a common enemy

(i)                      (which is why no change of significance is ever achieved by a government except during times of crisis; the flip side of this is that individuals who selfishly benefit from the status quo can always be expected to fight, with gusto, for their benefit from the way things are)

(d)                    [group selection (Google Search)] [index]

(46) Avoidance of punishment

(a)                    Lastly, individuals can be willing to display cooperative behavior if doing so means avoiding punishment (e.g., in human societies, as mediated by the police in human societies, or via banishment by friends for exhibiting a particularly selfish behavior)

(b)                    In evolutionary terms, however, punishment is some act by another that decreases an individual's Darwinian fitness; thus, cooperating to avoid a decline in Darwinian fitness is not necessarily an unselfish act

(c)                    Governments exist basically to foster altruism particularly by punishing selfish behavior either directly (e.g., sending people to prison) or indirectly (e.g., by rewarding selfless behavior such as by providing a break on taxes for contributing to charity or by paying people directly to work in the public's interest)

(d)                    [avoidance of punishment (Google Search)] [index]




(47) Vocabulary [index]

(a)                    Agonistic behavior

(b)                    Altruism

(c)                    Associative learning

(d)                    Avoidance of punishment

(e)                    Behavior

(f)                      Behavioral ecology

(g)                    Classical conditioning

(h)                    Coefficient of relatedness

(i)                      Cooperative behavior

(j)                      Courtship

(k)                    Critical period

(l)                      Dominance hierarchies

(m)                  Ethology

(n)                    FAP

(o)                    Fixed action patterns

(p)                    Foraging

(q)                    Generalists

(r)                     Group selection

(s)                     Habituation

(t)                      Home range

(u)                    Imprinting

(v)                    Inclusive fitness

(w)                  Insight learning

(x)                    Kinesis

(y)                    Kin selection

(z)                     Learning

(aa)                 Mating behavior

(bb)                Mating systems

(cc)                 Maturation

(dd)                Migration

(ee)                 Monogamous

(ff)                    Navigation

(gg)                 Observational learning

(hh)                 Operant conditioning

(ii)                     Optimal foraging

(jj)                    Orientation

(kk)                Parental investment

(ll)                     Piloting

(mm)             Play

(nn)                 Polyandry

(oo)                Polygamous

(pp)                Polygyny

(qq)                Promiscuous

(rr)                   Proximate causation

(ss)                  Reciprocal altruism

(tt)                    Releaser

(uu)                 Rhythmic behaviors

(vv)                 Search image

(ww)             Sign stimulus

(xx)                 Social behavior

(yy)                 Sociobiology

(zz)                  Specialists

(aaa)             Taxis

(bbb)            Territoriality

(ccc)             Ultimate causation