Storage, Ecology of

Caroline M. Pond , in Encyclopedia of Biodiversity (Second Edition), 2013

Hibernation

Hibernation and its hot weather equivalent, aestivation, are periods of inactivity and seclusion, usually accompanied by cooler than normal torso temperature, which enable animals to laissez passer through seasons when nutrient is scarce or inaccessible. Hibernation was amongst the outset physiological states in which adipose tissue of wild mammals, reptiles, and amphibians was studied thoroughly. Hibernators depend on their fat reserves while common cold and to fuel rewarming but reclaiming the stores is not every bit straightforward every bit it first seemed.

The enzymatic processes involved in fasting and starvation are essentially similar to those of slow exercise, but there is a critical difference: during do, the body is warm, ofttimes slightly warmer than when sedentary, only in hibernation, the trunk temperature is low, sometimes falling by 35   °C to close to 0   °C. The fatty acid composition of triacylglycerols is largely irrelevant to their role equally fuel when animals are warm, but it is crucial for their use during hibernation because enzymes do not work on solidified fats.

Experiments on captive chipmunks (Eutamias amoenus) and golden-mantle ground squirrels (Spermophilus lateralis) show that they enter hibernation more readily, remain cooler for longer and are improve able to survive long winters when plenty of polyunsaturated lipids are included in their nutrition during the weeks preceding hibernation than when they are fed on saturated fats of similar calorific value. Unsaturated fatty acids may lower the melting point of the triacylglycerols and the membrane phospholipids, enabling them to remain more fluid, and hence retain their proper affinity for carrier molecules and enzymes, at cooler temperatures. The chemical limerick of the storage lipids, together with other aspects of adipose tissue, is thus adapted to the physical weather condition and its role in whole-body metabolism (Frank and Storey, 1995; Frank et al., 2008).

Squirrels obtain many such unsaturated fatty acids from the seeds and other plant parts that they eat. Hibernation is an agile, physiologically controlled procedure, and especially in mammals, metabolic preparations can exist identified days or weeks before the animal really allows its body to cool. As the weather becomes cooler and the days shorten in autumn, squirrels, and other hibernatory rodents actively seek nuts and other foods that contain these lipids. The woodchuck or marmot (Marmota flaviventris) selectively retains linoleic acid (C18:2) before hibernation: the saturated fatty acids are released and oxidized by the muscles, liver, etc. while the creature is warm and active, but the polyunsaturates remain in the adipose tissue, for utilise when the trunk is cold. Like other mammals, the squirrels cannot add more than one double bond to most kinds of long chain fatty acids, then they depend on the increased availability in autumn of seeds rich in polyunsaturated lipids. Failure of a seed crop could foreclose successful hibernation and thus lead to death from cold or starvation, even if plenty of other foods were available. Analysis of biopsies of gonadal and inguinal adipose tissue from alpine marmots (Marmota marmota) throughout the year signal that selective release of sure fat acids allows active regulation of the composition of storage triacylglycerols (Cochet et al., 1999). The storage lipids remain fluid during deep hibernation and in the euthermic country by maintaining a high proportion of monunsaturates, both by retention of those from the nutrition and, if necessary, by synthesis.

Sleeping undisturbed in a cool, secluded place uses storage materials merely very slowly: small tortoises may lose less than five%, and rarely lose over fifteen%, of the torso mass during 5 months of hibernation at well-nigh v   °C. Many reptiles and amphibians sally from hibernation with a surprisingly large proportion of their lipid reserves even so remaining and regain what they lost over the winter in a few weeks of feeding. These stores are often of import for fuelling mating and egg production. The less lipid a hibernator uses during the winter, the more it has left to fuel breeding the following spring.

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Psychological Development: Ethological and Evolutionary Approaches

C. Trevarthen , in International Encyclopedia of the Social & Behavioral Sciences, 2001

Ethology identifies forms of action and perception that direct an animal's experiences in the world, and that mediare communication with animals of the same or unlike species. Beast motives, as products of evolution, are adaptive; that is, they generate behaviors that benefit the life of the animal. Innate actions, sensation, and feelings that initiate and guide learning, and are modified by learning, alter systematically during the lifecycle. Signaling behaviors enable animals to communicate interests, purposes, and feelings, and in highly social species, both innate motives and learning contribute to intricate forms of cooperation. Play behavior benefits evolution of skills, especially social skills, and human infants show complex intersubjective adaptations, which, in the imaginative, artistic play with companions, lead to acquisition of cultural meanings and language. Contemporary 'affective neuroscience' supports the view that intrinsic and largely innate motives and emotions of sub-cortical parts of the brain have a regulatory and formative role in the evolution and evolution of cognitive powers of consciousness, voluntary activity, thought, and memory. They guide human being cultural learning, contributing to the corking diversity of the human phenotype.

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A Role for Learning and Memory in the Expression of an Innate Behavior

Gabriela Rodríguez-Manzo , Ana Canseco-Alba , in Identification of Neural Markers Accompanying Memory, 2014

Introduction

Animal behavior can be broadly divided into instinctive or innate behaviors and learned behaviors. Instinctive behaviors are genetically determined; they are stereotyped (e.g., they are executed the same way every time); their expression requires no experience, meaning that they manifest without prior learning and consequently, retention is not of import for its display. Learned behaviors, past contrast, are acquired behaviors; animals are non born with them; their functioning is improved past feel and memory plays an important part in its expression. Learned beliefs can exist changed and supposedly instinctive behavior cannot. Withal, some innate behaviors may be modified (or modulated) through practice and feel, indicating that the classification of behaviors in either "innate" or "learned" might be excessive. Almost behaviors are a mix of the two, neither completely innate nor entirely learned (for review encounter Barnard, 2004; Bolhuis and Giraldeau, 2005). This is the case of sexual beliefs. In addition, instinctive and learned behaviors share a common neural substrate as evidenced by the fact that lesion of specific brain areas impairs both learned and instinctive behaviors. Thus, experimental destruction of the neocortex or the hippocampus in laboratory rats disrupts running through a maze without making errors and impairs the performance of sexual beliefs (Vanderwolf, 2003).

This chapter is aimed to review evidence for a role of learning and memory in diverse aspects of a behavior that is classified every bit instinctive: copulatory behavior.

Learning can exist defined every bit an adaptive change in behavior acquired by feel and the storage and recall of previous experiences describes the term memory (Kandel and Squire, 2009). Animals must be able to brand adjustments in their nervous systems in response to environmental stimuli that permit adaptive behavioral responses (Barnard, 2004). The ability of animals to arrange behaviorally in response to external stimuli is essential for survival. For a long time information technology was considered that this power was not involved in the brandish of instinctive, hard-wired behaviors which merely reflected the activation of developmentally programmed neural circuits that were non modifiable by the environment.

Sex activity is an innate highly rewarding behavior. According to Thorndike's Law of Effect (1911) a reward increases the frequency and intensity of a specific behavioral act that has resulted in a reward before (Schultz, 2006). The ability to learn about rewards is besides a crucial adaptive capacity and the ability to acquire and to remember what is learned is important to maintain adaptations (Barnard, 2004; Bolhuis and Giraldeau, 2005). The rewarding component of sexual activeness ensures its repetition contributing to guarantee species survival. The appropriate expression of an innate behavioral sequence frequently requires signals from the outside world (Burkhardt, 2005). A cardinal component of reward-related learning is the acquisition of conditioned associations betwixt rewards and ecology stimuli that predict or accompany those rewards. Such stimuli are called "conditioned stimuli" (Zellner and Ranaldi, 2010). The rewarding and reinforcing character of male person rat sexual behavior is evidenced by the fact that male person rats form a conditioned identify preference (CPP) for copulation (Ã…gmo and Berenfeld, 1990; Pfaus and Phillips, 1991; Tenk et al., 2009), demonstrating that the male rat is able to associate an environment with the rewarding sexual feel and is capable of recalling that association.

Sexual behavior is an innate behavior which expression is delayed until puberty. During this catamenia important hormonal variations underlie sexual maturation, which include hormonal, anatomical, and neurochemical changes in all sex-related tissues, including the brain (Schulz et al., 2009). The evolution of copulatory beliefs in rats depends on gonadal hormones, since its appearance is prevented by prepubertal castration (Larsson, 1967). Puberty-related encephalon changes trigger sexual behavior brandish in response to the adequate stimuli, thus male sexual behavior expression is regulated by brain circuits that are importantly shaped by sexual activity hormones such as testosterone and estrogen (Simerly, 2005). Although sexual development is adamant by steroid hormonal exposure, learning from social and sexual experiences at different stages contributes to the shaping of sexual function (Woodson, 2002). Sex hormones participate too in neuronal plasticity processes in the adult encephalon (Dugger et al., 2008); therefore, such neural plasticity might be 1 of the mechanisms underlying the learning processes associated to sexual behavior expression in developed individuals.

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Conceptual Problems in Cognitive Ethology

Colin Beer , in Advances in the Study of Behavior, 1992

A INTENTIONALITY

For cognitive ethology, intentionalistic readings of mind-laden words applied to animals should exist taken seriously and non set up bated as sociobiology stipulates. But by "intentionalistic" here, I mean more than the word usually connotes, and I need to make this explicit. I need to practice this because, as Dennett has said, "ethologists . . . persist in conflating the philosophical notion (Brentano's notion—the concept of aboutness, in a word) and the more than or less everyday notion of intentionality: the capacity to perform intentional deportment or frame intentions to act" Dennett, 1987, p. 271); and because the philosophical notion Dennett refers to is implicitly involved in much that cognitive ethologists say nigh the minds of animals.

However, this philosophical notion of intentionality appears to be hard for non-philosophers to go clear. A reader of a previous version of this article found my attempt at explication of intentionality incomprehensible. A recent book on motivation, past an ethologist, represents a special logical characteristic of intentionalistic idioms—their "referential opacity"—equally the "non-translatability of intentional terms into behavioural terms" (Colgan, 1989, p. 67), which is dead wrong. The philosophers do not brand things any easier when they vary the spelling between "intentionality" and "intensionality," sometimes using the difference to marker a distinction, and sometimes not. They also use the coupled terms "intension" and "extension" for the stardom between a term's conceptual meaning and what falls within its domain, as in the contrast between sense and reference, connotation and denotation (Cohen and Nagel, 1934). Moreover, intentionality, in the sense deriving from Brentano, continues to be a controversial matter for philosophers of mind (e.g., Dennett, 1969, 1987; Searle, 1980). In spite of these difficulties, I call up it necessary to include at least an outline of the concept here, for it could have serious implications for cognitive ethology. Other and fuller treatments can be plant in Dennett (1969, Chapter 2), Boden (1972, esp. pp. 47-49), and Lockery (1989, esp. pp. 121-129).

Brentano's (1874) concept of intentionality includes all mental states that are about or directed toward something or other (their content). And then in addition to intention, which is always intention that something or other come to laissez passer, intentionality includes knowing, which is always cognition nigh something or other, desiring, which is always desire for something or other, guessing, which is always guessing that or well-nigh something or other; likewise, recognizing, wishing, hoping, remembering, forgetting, doubting, and numerous other terms standing for what philosophers call propositional attitudes. As I pointed out earlier about perception, which is another example of intentionality, an intentional state is dissimilar a state like tiredness or lassitude in that y'all cannot, for example, but believe or desire purely and merely, empty of content. Brentano thought that only mental states could have this belongings of aboutness, that intentionality is "what distinguishes the mental of psychological from the concrete" (Chisholm, 1967, p. 201). If this were so, then to talk of animals as perceiving, recognizing, remembering, forgetting, and and then forth, would be tantamount to assuming them to have mentality. The exclusiveness of intentionalistic properties to mental states is one of the issues that philosophers continue to contend.

To support the thesis that intentionality is "the mark of the mental," philosophers have sought logical criteria that apply to sentences describing mental states merely non sentences describing physical phenomena. Among these criteria are referential opacity (Quine, 1960, p. 144) and existential generalization. Referential opacity obtains when substitution of coreferring terms in a judgement may not preserve its truth. Take the sentence "John believes that George Eliot wrote Middlemarch," and assume it to exist true. If yous substitute "Mary Ann (or Marian) Evans" for "George Eliot," the resulting sentence could exist faux, for John might not know that George Eliot and Mary Ann Evans are one and the same. Contrast this with "George Eliot wrote Middlemarch," which states a nonpsychological fact. Now the exchange makes no difference to the truth of the sentence.

For "George Eliot wrote Middlemarch" to be true, there must accept been a writer called George Eliot and a book chosen Middlemarch. Neither of these demand exist the instance for the judgement well-nigh John's conventionalities to exist true. This exemplifies the examination of existential generalization. As a rule, sentences apropos nonpsychological facts (extensional sentences) presume the existence of what they refer to; intentional sentences need not.

Although these and related logical criteria more often than not differentiate sentences about intentional states from sentences about concrete facts, in that location are exceptions (see Chisholm, 1967; Kneale and Prior, 1968), particularly cases of nonintentional description that sideslip by the logical tests for intentionality (e.g., the nonintentional sentence "three is the foursquare root of ix" appears to be referentially opaque; endeavour substituting "the number of the planets" for "nine"). Philosophers are still seeking ways of refining the logical rules then that they pick out all and simply statements about intentional states. In the meantime, nosotros can ask whether and how these logical tests utilize to ascriptions of intentional states to animals.

The answer seems to be that it is hundred-to-one whether there can ever be a straightforward awarding of whatever of the logical tests for intentional ascription in fauna cases. Take the possibility that an animal might believe in or desire something that does not exist. For this to exist judged seriously, there must surely be some observable or empirical consequences. It is hard to remember of any cases for which what might be construed as appreciable consequences of such a conventionalities or desire cannot be interpreted in simpler terms, making no appeal to intentionality. For example, when a gull or a fish reacts to its mirror image as though it believed it were faced past another of its kind, an ethologist can fall back on the classical story of how sign stimuli elicit patterns of ambitious or courtship behavior. Even the bees that Gould (Gould and Gould, 1986) tricked into passing fake information to their hive mates tin be accommodated past a causal business relationship needing no importation of intentionality. Millikan has viewed the bee dancing every bit referentially opaque: "Von Frisch knew what bee dances are about, but it is unlikely that bees do. Bees but react to the bee dances accordingly" (Millikan, 1984, p. xiii). But this manner of deploying referential opacity is hardly a exam for intentional ascription comparable to a homo case. If it were, we could credit a clock with intentionality, considering we know what the position of the hands on the clock face are about, but the clock does not. Unless the bees have alternative ways of representing the same state of affairs and, hence, are in a position to be acquainted with this state of affairs under one representation but not under another, they cannot be capable of being in referentially opaque states in the way that nosotros are. Indeed, when Millikan says that the bees merely react appropriately, she admits equally much.

We accept culling mental representations of the same things because nosotros know them by different signs, different names, different descriptions. Then we tin can refer to the same heavenly body as "the forenoon star," "the evening star," or "the planet Venus"; whether we know that these alternatives refer to the aforementioned affair will affect how they enter into our beliefs. Because animals, with the possible exception of some peculiarly trained instances, lack anything like our cerebral manifold of names and descriptions, trivial if annihilation of what logically applies to intentional ascription in the human being case carries straightforwardly over to intentional ascription in animal cases.

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Of Course Animals Are Creative

Robert Epstein , Dean Keith Simonton , in Animal Creativity and Innovation, 2015

Generativity Theory

Whether novel fauna behavior has value in the same sense that novel human behavior has value is debatable, just every bit the significance of computer-generated art and music is debatable, or fifty-fifty the significance, for that matter, of any estimator behavior that seems to mimic human intelligence (Epstein, Roberts, & Beber, 2008; Turing, 1950). But the fact that animals exercise new things is indisputable. Where does novel behavior come from, and is it possible that the same basic processes are responsible for the novel beliefs of both animals and people?

In the early 1980s, I proposed a theory to endeavour to explain the emergence of novel beliefs in multiple species. According to generativity theory, (i) new behavior emerges equally previously established repertoires of beliefs become interconnected over time, and (ii) the interconnection process is both orderly and predictable. Combinatorial theories of inventiveness have been around for a long time (consider Hull, 1935; Poincaré, 1946; Rothenberg, 1971); the main thing I added to the moving-picture show was an assertion of orderliness, likewise every bit a mode of quantifying and measuring this orderliness. Specifically, I speculated that multiple processes operate simultaneously on the probabilities of multiple behaviors, and, by implication, on their underlying neurological systems.

This is a sensible and, in some respects, uncomplicated theory. It is also predictive and parsimonious. Most important for present purposes, generativity theory explains why novel behavior—a small portion of which volition inevitably have value in specific situations—is ubiquitous in the beast kingdom. The orderly and rapid interconnection of repertoires of beliefs is adaptive; it is near every bit adaptive a process equally nature could perchance devise considering information technology guarantees that all previously established behaviors, whether learned or programmed by genes, will be available to tackle new challenges as they ascend moment to moment in time in the environment. Generativity theory shows how this process works without making any assumptions about speculative cerebral mechanisms—or denying their existence.

The same mechanisms underlie the emergence of novel behavior in both animals and people because novelty-yielding behavioral variability has value, just every bit novelty-yielding genotypic and phenotypic variability accept value in development. Species survive and sometime diverge because of phenotypic variants. The range of variability in traits yielded by sexual reproduction is and so large that genuinely new traits inevitably sally in every generation—a miracle that helps to protect a species from extinction when ecology conditions change and that ensures that superior traits and even superior species will ultimately triumph over time. Similarly, novelty-yielding behavioral variability in an individual organism helps to guarantee that that individual's behavior volition be effective under changing atmospheric condition, occasionally fifty-fifty producing behavior and so new that it tin can alter the organism'due south environment in pregnant means.

These two types of variability might fifty-fifty overlap. Genes might occasionally produce individual organisms so creative that they find effective new means to compete against other species for express resource, eventually dislodging those species from the cistron pool.

2 ubiquitous situations in the natural surround assure that behavioral competition occurs nearly continuously. First, the existent world surrounds united states constantly with multiple, novel, and vague stimuli which ready multiple behaviors in movement. Second, when behavior is ineffective—which it is in pocket-size ways hundreds of times a day—a process called "resurgence" occurs: previously established behaviors that were constructive under conditions similar to the electric current ones are activated (Epstein, 1983, 1985b, 1996a; cf. Epstein & Skinner, 1980). Generative processes thus assure not only that behavior volition vary just that information technology will vary in ways that are especially probable to exist adaptive in the current situation. In this sense the variability that occurs in individual organisms is even more than adaptive than the variability that drives the evolution of species; the latter is blind, merely the quondam guarantees the emergence of new behavior that is specifically relevant to the properties of the new situation. Generativity theory too has immediate practical value, considering when you place variables and parameters that contribute to the emergence of novel behavior, you lot can manipulate those variables and parameters for useful ends (Epstein, 1996b, 2000, 2011; Epstein, Kaminaka, Phan, & Uda, 2013; Epstein & Phan, 2012; Epstein, Schmidt, & Warfel, 2008).

To explore the potential power of generative contest, I expressed the theory in formal terms and modeled it with a computer algorithm. Representing but four long-studied behavioral processes in a series of equations I called "transformation functions" (Effigy thirteen.1), I was able to model a number of creative performances I had studied in laboratory settings with both pigeons and people, such as the manner in which people solved Maier'southward (1931) archetype "2-cord problem" (Figure 13.2). Running simultaneously in a "state" algorithm, the transformation functions generate a "probability profile," in which overlapping probability curves show how behavior changes over time, producing novel beliefs almost continuously equally other behaviors go interconnected over time (Figure xiii.iii). I also devised a way to prove the orderliness in the novel performance of a single subject area using a graphical technique that generates a "frequency contour." The frequency profile yields overlapping curves that look very much like probability curves (Figure 13.4).

Figure 13.i. The transformation functions of generativity theory.

According to generativity theory, multiple behavioral processes operate simultaneously on the probabilities of multiple behaviors. In one possible instantiation of the theory, iv basic behavioral processes are represented (above). y n is the probability of behavior y at bike northward of the algorithm, y′ north is the probability of behavior y′ at cycle n of the algorithm, ε is a constant for extinction (it determines the charge per unit at which the probability of behavior y decreases over cycles of the algorithm), α is a abiding for reinforcement (it determines the charge per unit at which the probability of behavior y increases over cycles of the algorithm as a result of certain environmental events), and λ yy′ is the constant of interaction between behaviors y and y′.

Figure 13.2. Maier's (1931) Two-string problem.

Subjects are instructed to tie the ii ends of the strings together, simply they quickly larn that they can't reach both strings at one time. They acquire this past pulling one cord toward the other and reaching. Most people then endeavour pulling the second string toward the first, which makes footling sense. When provided with a long heavy object (#v in inset), a subject is highly likely to use information technology to extend his or her reach, only the object that is provided is not long enough to achieve the other string. When provided with a brusk heavy object (#1), a subject is much more likely to solve the trouble, which requires tying the object, brusque or tall, to ane string and swinging it, so pulling the other string toward the swinging cord and catching information technology when it comes near. Appropriately, the problem is sometimes chosen "the pendulum trouble." Provided with a long object, if a subject is able to solve the trouble at all, automated chaining is usually involved. The person ties the long object to the end of a string and so pulls the object toward the second string; this is one fashion of using the object to extend one's reach. When that fails, the subject field ofttimes lets get of the object, which causes the attached string to swing in a pendulum motion. The solution follows speedily. Objects of intermediate lengths produce predictable outcomes according to those lengths.

Figure xiii.3. Probability profile for Maier's (1931) 2-string problem.

A probability contour generated by the transformation functions shown in Figure 13.1, generated for five behaviors relevant to Maier's (1931) two-string problem. The abscissa is labeled "ticks," which are cycles of the computer algorithm, each a scalable moment of unspecified duration. The contour was generated with parameters for a brusque object (#1 in Effigy 13.2), which generally produced rapid solutions to the problem and no irrelevant reaching. Annotation that pulling one string toward the other decreases steadily in probability and that other behaviors increase in probability in an orderly sequence. Tying the object to the cord makes swinging more than likely, which, in turn, makes connecting the strings more likely. The figurer model that generates the curve uses discrete state methodology, running a set of initial probabilities through all 4 equations to generate a new set of probabilities, then running those through the equations once more, and then on.

Figure 13.4. Predicting individual beliefs moment to moment in time.

(A) This probability profile, produced by the transformation functions of the generativity model, predicts the behavior of a human subject on a touch-screen task. The subject has been instructed to movement a spot across the screen into a goal expanse. Borer three patches on the screen (B1, B2, and B3) will move the spot in various directions and at varying speeds; tapping a fourth patch (B4) has no effect. The model predicts that the subject will begin borer B1, then gradually shift to B2, then gradually shift to B3, with responses alternating among the iii choices along the way (where the curves overlap). It also predicts that toward the end of the session, the subject field volition brainstorm tapping B4, even though doing and then has no outcome. (B) This frequency profile shows actual information obtained from one subject area (S58) during a 5   min session. The pattern of responding is predicted well by the probability contour, including the shift to B4.

This methodology proved effective in predicting novel performances in both pigeons and people in laboratory performances. The differences between the 2 species seem largely to be parametric. The general principles—that new behavior emerges as erstwhile behaviors merge and that multiple behavioral processes operate simultaneously on the probabilities of different behaviors—seem valid for both species and perchance for many others as well.

The formal representation and quantification of the creative process in private organisms is no small feat, simply with few exceptions the existing literature on animal creativity and trouble solving shows little or no sensation of any of the main implications of generativity theory and its supporting research. Instead researchers are (and I say this respectfully) wasting their time debating about what animals may or may not "understand" regarding their performances, equally if that data—were it even possible to decide with any caste of confidence, which it is not—would add anything important to our own "understanding" of the performances.

When y'all speculate about what an animal "understands" regarding an arrangement of stimuli or the manner in which information technology solved a trouble, you are simply talking most more than behavior. You are asking whether the creature can not only solve the problem but tin can also country a principle or visualize a causal menstruation diagram or perhaps even perform mathematical calculations. The trouble hither—which has existed as long equally comparative psychology has existed—is that as long as you can excogitate of a fashion for the creature to accept solved the trouble without engaging in these so-called "high-level" cognitive manipulations, you must make the parsimonious supposition that the animal is not doing so.

Even if we were able to prove definitely that an animate being had formulated the (human) verbal equivalent of a formal principle, I accept little doubt that the processes giving ascension to the emergence of such a principle—once more, this is only more beliefs, subsequently all—would be like if not identical to the processes that led to the emergence of the problem-solving operation itself. In footnote v of an early newspaper I published on the principle of resurgence, I explained how generative processes—in particular, the process of resurgence itself—could business relationship for my formulation of the formal principle of resurgence itself (Epstein, 1985b, p. 151; besides see Epstein, 1996a, p. 145). A argument of a relationship among variables—in other words, "reasoning"—is, first and foremost, a argument; again, information technology is merely more behavior, presumably amenable to the same sort of analysis that tin be applied to all behavior.

Without exception, all of the studies of animal trouble solving or creativity I have seen in recent years lend themselves to a rigorous analysis using the tools of generativity theory. Video recordings can exist examined frame past frame, coded, and represented past frequency profiles, which instantly reveal orderliness which is almost entirely invisible to the naked eye. Instead, researchers are all the same relying on crude verbal descriptions of the performances, or, at most, crude tabulations of "per centum correct" and other data aggregated across organisms or trials. Trouble situations, which past definition, are divisional in specific means, can easily be represented in formal terms using transformation functions, and those functions can then be used to model and predict individual performances. In short, the orderliness in animal inventiveness can be quantified and studied rigorously using advanced tools of the natural sciences.

Whether generativity theory is correct in its particulars is beside the point. The predictive power of this type of theory is and so corking that some form of it almost certainly must be correct. Almost certainly, multiple processes must be acting simultaneously on the probabilities of multiple behaviors and their counterparts in the nervous system, and the cyberspace issue generates a wide range of behavior continuously in time—everything from mundane preparation to profound "insight."

Meanwhile, many researchers in psychology and biology who are rightfully fascinated by some of the extraordinary man-like capabilities of crows, chimpanzees, and other animals, are barking up the incorrect tree. Similar the naturalists of the 1800s, they go on to anthropomorphize, insinuating that human-like performances past animals are interesting merely if an animal'south cognitive earth is like a human's.

Meanwhile, I wonder, as many have before me, why nosotros continue to bother speculating nigh the cognitive world of humans; I have long seen this equally a dilemma in which our consciousness interferes with our scientific objectivity (Epstein, 1982, 2008).

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Emotion, Evolution of

Elena Walsh , Paul E. Griffiths , in International Encyclopedia of the Social & Behavioral Sciences (2nd Edition), 2015

The Ethological Tradition

The founders of ethology saw themselves as the direct heirs of Darwin'due south piece of work on mental evolution (Lorenz, 1965). They retained Darwin'southward principles, merely reinterpreted them to fit the theory of evolution as information technology emerged in the 1930s in the 'modern synthesis' of Darwinism and Mendelian genetics. The principle of serviceable associated habits was transformed into the ethological concepts of 'ritualization' and 'derived activity' (Tinbergen, 1952). Derived activities are behaviors that originally evolved for i purpose only were later selected for another purpose. Ritualized behaviors are derived activities that originally evolved to fulfill some practical function but which were subsequently selected to function every bit signals. Thus, although piloerection in fearfulness and rage does little to make a human appear larger to an opponent, it does communicate their emotional state. Derived activities require a distinctive, 2-phase form of evolutionary explanation. They cannot exist understood purely in terms of the office they currently perform and the pick pressures that currently maintain them in the population. This is particularly obvious in the case of signals. Piloerection is non intrinsically better as a signal of fearfulness than smiling or laughing. This item behavior was selected as a signal only because information technology was already associated with fear in the distant past. It was not associated with fear because information technology was a signal of fright, but considering it fabricated the animal appear larger.

The concept of ritualization immune ethology to reconstruct Darwin's principle of serviceable associated habits while fugitive his commitment to the inheritance of caused characteristics. Darwin's descriptions of the psychological rewards that led to the reinforcement of emotional behaviors are equally plausible every bit descriptions of the original selective advantage of those behaviors. Darwin'southward other two principles were similarly open to reinterpretation. The principle of antithesis was explained by the selective value of unambiguous signals: it may exist important for an animate being to betoken clearly that it is not aggressive. Hence there can be pick of behaviors but because they look unlike from the behaviors that bespeak aggression. The principle of direct activity was transformed into the ethological concept of displacement action. Early on ethologists shared Darwin'due south view that instinctive motivations crusade a build up of mental energy that must be released in some behavior or other. Deportation activities "serve a role every bit outlets, through a rubber valve, of unsafe surplus impulses" (Tinbergen, 1952: p. 23). An example commonly given is that of an angry cat that is unwilling to assault and begins to wash itself. This wholesale reinterpretation of Darwin's iii principles worked and so smoothly and immune the retention of so much of the detail of Darwin's work that the early on ethologists seemed almost unaware of the differences between Darwin's theory and their own (Lorenz, 1965).

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Emotions, Evolution of

P.Due east. Griffiths , in International Encyclopedia of the Social & Behavioral Sciences, 2001

ii The Ethological Tradition

The founders of ethology saw themselves as the direct heirs of Darwin'south piece of work on mental evolution (Lorenz 1965). Their account of the evolution of emotion retains Darwin's principles, but reinterprets them to fit the theory of evolution equally it emerged in the 1930s in the 'modern synthesis' of Darwinism and Mendelian genetics. The principle of serviceable associated habits is transformed into the ethological concepts of 'ritualisation' and 'derived activity' (Tinbergen 1952). Derived activities are behaviors that originally evolved for one purpose but were afterward selected for another purpose. Ritualized behaviors are derived activities that originally evolved to fulfill some practical role merely which were later selected to function as signals. Thus, although piloerection in fear and rage does non brand a human being appear larger to an opponent, information technology does communicate his or her emotional state. Derived activities crave a special pattern of evolutionary caption. They cannot exist understood purely in terms of the role they currently perform and the pick pressures that currently maintain them in the population. This is peculiarly obvious in the case of signals. Piloerection is not intrinsically better as a signal of fear than smiling or laughing. This item behavior was selected as a bespeak only considering it was already associated with sure emotional states in the afar past. It was associated with those states, not because information technology was a signal, only because information technology made the animal appear larger.

The concept of ritualization allowed ethology to reconstruct Darwin's principle of serviceable associated habits whilst fugitive his commitment to the inheritance of caused characteristics. Darwin's descriptions of the psychological rewards that led to the reinforcement of emotional behaviors are as plausible as descriptions of the original selective advantage of those behaviors. Darwin'southward other 2 principles are equally open up to reinterpretation. The principle of antithesis is explained by the selective value of unambiguous signals. It is as important for a domestic dog to betoken that it wants to avoid conflict as for it to point aggression. Hence at that place tin can exist selection of behaviors merely because they expect different from the behaviors that signal assailment. The principle of straight action was transformed into the ethological concept of a displacement activity. Early on ethologists shared Darwin's view that instinctive motivations cause a build up of mental free energy that must exist released in some beliefs or other. An example commonly given is that of an angry true cat that is unwilling to attack and begins to launder itself. Niko Tinbergen remarks: 'I think it is probable that displacements practice serve a function as outlets, through a safety valve, of dangerous surplus impulses' (Tinbergen 1952, p. 23). This wholesale reinterpretation of Darwin's iii principles works so smoothly and allows the retention of so much of the detail of Darwin's work that the early ethologists seem nigh unaware of the differences between Darwin'due south theory and their ain (Lorenz 1965).

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Crying

B.Yard. Lester , L.L. LaGasse , in Encyclopedia of Infant and Early Childhood Evolution, 2008

Distress Calls Among Mammals

Ethologists, who report animal beliefs, telephone call crying a proximity-promoting and -maintaining signal, that is, it encourages the female parent to stay with and sooth her babe. In that location are many species that can immediately locomote or in some mode follow their mothers at nativity. But relative to other species, human infants have a long period of dependency. Many species including cats, bats, elephants, seals, and reindeer, and of course chimps, utilise a distress vocalization or cry to indicate the mother when the infant is isolated, hungry, or cold. The prosodic features of speech described above are the major component of the vocalization of most mammals. Researchers have analyzed acoustic patterns and shown that nonhuman distress vocalizations show similar patterns to homo infant cries. Infants depend on song communication to signal distress or otherwise summon the mother close. Crying turns out to be an exceptionally effective survival mechanism. It is an information transmission system that sends melancholia messages, for example, hunger, pain, and demand for attention. Another example of how this remarkable system works comes from opera.

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The Evolution of Behavioral Phenotypes: Lessons Learned from Divergent Spider Populations

Susan Eastward. Riechert , in Advances in the Study of Beliefs, 1993

D CORRELATED TRAITS

Many contemporary studies of beast beliefs attempt to explain behavioral development by focusing on one functional category of behavior at a time (due east.g., foraging beliefs, agonistic behavior, mating beliefs) and measuring or inferring its contribution to the fitness of individuals. Notwithstanding, genes may influence many different characters, the manifold effects of a single gene existence referred to as pleiotropism. Oftentimes there is resulting correlation between morphology and beliefs, equally in the archetype work by Sturtevant (1915) in which Drosophila heart color was constitute to be genetically correlated with level of sexual activity. There also may be linkage betwixt loci that are closely positioned on the same chromosomes such that they segregate together.

In the calorie-free of these genetic phenomena, Dobzhansky (1956) stated that "a trait has no adaptive significance in isolation from the whole pattern that the organism exhibits." Huntingford provides a nice example of the need to consider the suite of behaviors rather than single traits. She establish that stickleback territorial aggression toward conspecifics during the convenance season covaries in the private with "boldness" toward predators in the nonbreeding flavour. Thus, individual fish that are well-nigh successful in obtaining breeding sites are apparently likewise most vulnerable to predation. Huntingford (1976) hypothesized that territorial aggression and disrespect toward predators shared some common factors(due south) within the fish. More recently, intraindividual covariance betwixt functionally different traits in some vertebrate species has been specifically attributed to the activeness of hormones such every bit testosterone. In several bird species, males with college testosterone titers show enhanced levels of territorial aggression and courtship but lower levels of parenting than other males (e.g., pied flycatchers: Silverin, 1980; firm sparrows: Hegner and Wingfield, 1987).

My colleagues, John Maynard Smith and Ann Hedrick, and I accept investigated the possibility that iv seemingly unrelated fitness-related behaviors described in this section are under the aforementioned genetic influence. Riechert and Maynard Smith (1989) found that individual spiders within a population that tended to win territorial disputes likewise demanded larger territories. Spiders that exhibited longer latencies to assail prey as well had longer latencies to return to a foraging mode following a predatory cue (Riechert and Hedrick, 1990). Nosotros have also established a link between the territorial behaviors and the attack and fear behaviors. A spider that showed a shorter latency to return to foraging following a predatory cue had a high probability of winning a territorial dispute against a spider that was equal in weight just had a longer latency to render to foraging. (Probability of being less fearful toward predatory cue and winning territorial dispute: NM feral individuals, .80; NM F2 generation lab.-reared .89; AZ feral individuals, .lxx; AZ F2 generation lab.-reared, .84.)

Maynard Smith and Riechert (1984) developed a genetic model to explicate the betwixt-population differences noted at the time in territorial and agonistic behavior. The model assumed that genes controlled two antagonistic hormones, 1 governing levels of fear and the other levels of aggression. If nosotros assume that at that place is directional authorization between the two populations in the genes that control levels of assailment and fear, as shown in Fig. 7, then the highly aggressive phenotype that results from crosses betwixt the ii populations is explained. Through written report of the backcrosses and F2 generation hybrids betwixt AZ riparian spiders and NM grassland spiders we take further found that the assailment components of agonistic and territory size behaviors are controlled by genes on the sex chromosomes, while the fearfulness component is controlled by multiple genes on the autosomes (Riechert and Maynard Smith, 1989).

Fig. 7. Proposed directional say-so mechanism of inheritance of aggression and fearfulness components of Agelenopsis aperta beliefs. Phenotypes and genotypes for AZ riparian and NM grassland individuals are shown, as well as the predicted outcomes of betwixt-population matings.

 (From Maynard Smith and Riechert, 1984.)

Finally, nosotros conclude from these studies that all iv traits probably stand for pleiotropic effects of the same genes, rather than linkage among chromosomes. The genetic system is simply too complex and involves besides many genes for linkage to be a likely cause. Nosotros besides wait that these genes command the production of given hormones or threshold responses for their release.

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Copying and Mate Choice

LEE A. DUGATKIN , in Social Learning in Animals, 1996

The Game Theory Approach

Game theory models of beast behavior await at the evolution of strategies, when the fettle of one private is affected not but by its own actions, but by the actions of others (Maynard Smith, 1982). The simplest possible game involving mate-choice copying involves a copying strategy (copy the mate pick of other females) and a chooser strategy (independent cess of male quality) competing with one another in an infinitely large population. Pruett-Jones (1992) models such a game in which all individuals have a baseline fettle, W. Mate assessment has a toll, one thousand, but those who assess males receive some benefit, f. Analysis of this game shows that when f > grand, copying and choosing coexist in a population at frequencies k/f and i – (thousand/f), respectively.

Losey, Stanton, Telecky, & Tyler (1986) built a computer simulation that examined copying and choosing strategies in a structured environment which imitation a lek breeding area. The distribution of male genetic quality was set in whatever given run of the simulation (but could be changed across runs). A second distribution, that of female choosing ability, was as well preset. This distribution determined the probability that a female using the choosing strategy mated with a male of a given genetic quality. Females using the copying strategy were immune a stock-still number of "peeks" at the choice made by other females (copiers and choosers alike). If a copier had not observed any males mate she either selected just as chooser would (the "smart copier" model) or she selected males randomly (the "impaired" copier model); withal, if she had seen males mate, she but chose the male that she had seen mate the greatest number of times.

For most of the parameter space explored, Losey et al. (1986) found that a mixture of copiers and choosers, with copiers in the minority, was the "evolutionarily stable stage." This frequency dependence was due to copiers copying each other rather than choosers, as the frequency of copying surmounted a critical threshold. Their model generates iv other testable predictions about female mate-choice copying in lekking species: (1) copying fares best when choosers are able to correctly identify high quality males, (2) copiers should take an reward when contest between young plays a pocket-sized role, (3) copiers should take an advantage when fecundity is depression, and (4) copying should fair well when females visit leks for either long periods of time, on numerous occasions, or both.

Using a simulation technique similar to Losey et al. (1986), Dugatkin and Hoglund (1995) examined mate-choice copying, when the tendency to copy is a continuous variable, and when female person reproduction is time constrained (i.e., if females spent too much time assessing males, they suffered reduced fecundity). In this model, females moved from territory to territory in a random fashion, and only a single female person was assumed to be on a given male'due south territory at once. All females assessed "male quality" and used a rule that instructed them to mate with a male person if its perceived genetic quality was greater than some value (V),simply female assessment was bailiwick to some error. Dugatkin and Hoglund assumed that a copier's, but non a noncopier's, perceived value of male quality is affected by the selection other females make. Females take a limited breeding season and consequently the more than time units that pass before they mate, the lower their fitness at mating. The tendency to re-create can thus have contradictory furnishings. While copiers may avoid the costs associated with mating tardily, because all females make errors in assessing a male's quality, copiers may copy others who themselves are making "bad" choices—i.east., choosing males whose real quality is less than 5.

Results point that the only variable that had articulate-cut interpretable effects on the evolution of copying was V, the minimum perceived male quality for female mating (Dugatkin & Hoglund, 1995). As V increased, the probability that a copying strategy invaded the population increased. Increasing V probably favored copying, because as V increased fewer males had the perceived quality score required past females. When any female, notwithstanding, did mate, copiers increased their quality assessment of that male, making it more than likely that they would mate early in the sampling process and thus avoid the costs of late mating. Different the case of Losey et al.'due south (1986) model, the copying strategy in Dugatkin and Hoglund (1995), for the most part, did better as the frequency of copiers increased (except for loftier values of V and many copiers). The positive frequency-dependent event found was tied to the cost of mating belatedly in the breeding season. Basically, some copiers (besides as non–copiers) mated early in the season. The more copiers at that place were, the more than early matings copiers saw, and the more than likely they were to lift their assessment of male qualities and thus to mate before themselves—hence the positive frequency dependence.

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