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Change topic • Enough on Proximate causes, interesting and a lot of work. • Basics: Stimulus: effector: in between: behavior • What we will look at now is the behaviors themselves! Will drift back to proximate causes but main focus will be behaviors • First distinguish between two major categories of behavior: Innate (instinctive) vs Learned Innate behaviors • Used to make sharp distinction between innate and learned behaviors. • Innate: as instinctive indicates, behavior that is “born with you” and can manifest itself without learning it. • First proposed by Konrad Lorenz early 1900’s Fixed action pattern • One of the standbys of innate behavior was the “fixed action pattern” • Defined as a behavior, once stimulated to start will continue to its end, even if the original stimulus (innate releasing mechanism)removed. • Greylag goose of Lorenz. Used to be the Classic example! • Now, hardly heard of!! Innate behavior • Why?? • Lot of controversy over this concept and in fact can’t even be found in many new books! • Here we touch on it for both historical reasons and just in case it becomes in fashion again! • Fairly stereotypic behaviors within species and across species. • Would manifest itself in isolation, no outside environmental influence, or at least we think so! Innate Learning • Many argued that can’t isolate from all possible environmental influence, so can never be sure…. • A lot of circular arguments made: graduate students trying to make a name for themselves! • U.S. vs Europe (reductionist vs holistic) • Yet we do recognize that behavior has a genetic basis! Innate behaviors • And that where it stands to day, so far… most accept that behavior has genetic basis (or at least control!!), just arguments as to how much!! • What are you to do? How to view innate vs learning • Best visual model is: all born with a framework (skeleton) of behaviors or behavioral capabilities: a lot due to anatomy and physiology: a dog may want to climb a tree but it will never learn how to do it! • This defines the range of possibilities. The actual behavioral repertoire (fleshing out of skeleton) individual develops depends on experiences (learning) it has. Phenotypic plasticity • Some equate it to “phenotypic plasticity” • Ability to change phenotype under different environmental conditions. • Example Bryozoans (spines in presence of predators but not in absence) • Example Locust (solitary vs migratory types: different structure AND different behaviors!) • Maybe Lamarck was right!!! Behavioral plasticity • Behaviors are more changeable than anatomy or physiology! • Although the behavior itself may not be genetically transmitted, the ability to do so is AND as we will see, can be transmitted socially. • Still all quite confusing!! Mental gymnastics!! Blah Blah Blah…. • What we do know for sure is that behaviors CAN change with experience, animals can learn. • So now looking at those modification from experience: learned behaviors (Chap. 4) Learned Behaviors What is learning • First must address the question of what do we mean when we talk about learning? • Or definition of learning: • A relatively permanent change in behavior as a result of experience. • One could argue: where did that original behavior come from?? Innate, Genetic? • Lets not go down that road!!!! Learned behaviors (chap 4) • Best to start with the behavior that enables many others: Learning behavior • Identified earlier as a source of gaining behaviors • Is a behavior in itself! Individual learning • Make a distinction between how an individual learns from experience and social learning, learning from other individuals. • Here we deal with individual learning • Three interrelated question: • 1) How do animals learn • 2) Why do animals learn • 3) What do animals learn How do animals learn?? • Psychologists: processes underlying learning. • Three types of experiences that lead to learning • 1) single stimulus • 2) Stimulus-stimulus • 3) response-reinforcer Single Stimulus • • • • Single stimulus given an animal: E.g. blue stick to a rat in a cage. Rats will notice because of novelty. If consistently notice it: call sensitization to the stimulus • If eventually ignores it: called habituation. • Two possible learned responses to a stimuls Single Stimulus • Sensitization is good If stimulus is important and may help animal associate this stimulus with other cues. • Habituation is good in that it helps reduce the background “noise” (bombarded with stimuli constantly) and enable response only to those of importance • Can be disadvantage if become habituated to a cue that later becomes important! Predation e.g. Stimulus-stimulus • Here we see how an animal related TWO stimuli • Example: pair blue stick with odor of a cat. • Cat reacts to odor initially but… • Eventually will hide just when blue stick is spotted: associates blue stick with cat.. • Pavlovian or classical conditioning. Terms…Terms..Terms! • Things getting complicated so need terms to keep it straight! • 1) First stimulus: Conditioned stimulus (in itself, does not elicit a response • 2) Second stimulus: Unconditioned stimulus (this is one that would normally elicit a response, even without training. • 3) Conditioned response: This is reaction of animal to conditioned stimulus AFTER it has associated it with unconditioned stimulus Terms…terms…and more terms! • A few more terms associated with this. • 4) Appetitive stimulus: stimulus that is considered positive, pleasant, or rewarding, e.g. food, potential mate, safe haven, etc. • 5) Aversive stimulus: Opposite – unpleasant, e.g. shock, noxious odors, etc. • 6) Excitatory conditioning: positive relationship – first stimulus leads to or predicts the occurrence of the second And more….. • Inhibitory conditioning: If first predicts second event will not occur: Huh? • E.g. if blue stick becomes associated with NOT getting food when rat would normally get it. • No blue stick, will get food. • Blue stick, no food. • Can add second and even 3rd and 4th conditioning stimuli!!! What does this all mean!!??? • Basically it means that animal can associate one stimulus with others AND to predictable outcomes (presence of cat, getting food or not….) • Through this association it “learns” how to modify its behavior • So the ability to associate stimuli with outcomes is important part of learning behavior. here Response-Reinforcer • This associate ability becomes more evident in third type of experience. • Commonly called: Instrumental conditioning or Operant or Goal-directed learning. • Response of animal is reinforced by a reward or punishment How different from Pavlovian? • Animal takes some action first, stimulus comes second. Associates outcome with stimulus rather than stimulus with outcome • Classic is rat pushing lever to get food. • Can get complicated! • Law of effect: If response in presence of stimulus = satisfying event: association between stimulus and response strengthened. And visa versa! • Skinner and “Skinner box” biggies in this area Summary on how they learn • Why is this all important? • We know animals (us!) learn from experience. What we need to know is how is that accomplished? • This then helps us understand how, why, and when certain behaviors may occur Summary • Results so far: A feedback system where, • 1) animal does something and gets a result and it remembers this • 2) something happens to animal and it remembers circumstances (stimuli) under which it happened • So two biggies: association AND Memory! • How memory works, is anyone’s guess!! Memory • Without it learning is impossible • Still don’t know exactly how we physically store info, some three dimension electrophysical process!!??/ • What we do know is: • Memory is selective! (extinction curves) • Memory capabilities vary intra and interspecifically: some better than others Memory • Brain-memory-learning is the main genetic link between genes and behavior. Why animals learn • Ok, some idea of learning process: carrot and stick! • Why do they learn? • Simple answer is that it has a “survival advantage” • But not as simple as that or is it?? Why learn • The fact that memory is selective can provide clues: • - Receive thousands/millions of stimuli • - Make thousands of associations • - Some we remember/others we forget • - Some we remember longer than others • Reflects importance of the associations with environment Why animals learn? • Behavior is interface of animal with its environment: a changing environment! • With this interface, animal needs to capture energy and avoid loosing it! • In capturing energy (food), learning important way of increasing foraging efficiency – reduce foraging time Why animals learn • In capturing “new energy” (reproduction), learning increases chance you will participate. (not just trees releasing pollen!). • In avoiding loosing energy, it is helpful to learn what are safe vs dangerous areas! So why do they learn? • Helps maintain the energy capturing machine under environmental conditions. • Indeed if they do not learn, they are dead meat!!! What do they learn? • Ok, out of those millions of stimuli and associations, which, if any, are important to learn about?? • Obviously rest of book is on the different behaviors that have developed based on stimuli received! • Here is a general treatment of the major categories, which we will then look in detail later. What do they learn? • List of things important to their survival! • 1) learning where home is located! • What this involves is all the various behaviors we will cover in Chapter 13! • As mentioned, behaviors are interface of animal with its environment Call Home!!! • Animals live in the environment and so it is important to learn about it. • Bird and other migrations: learning migratory routes • Habitat selection: learning what is the appropriate habitat to live in!! • Will see this is one of THE most important behaviors to learn for many species! What else? • 2) Learning about your mate!!! • Obviously important for the capture of future energy for the species!! • Two chapters devoted to this!! (6 & 7) • Need I say more?? Next on the list… • 3) Learning about familiar relationships (Chap 8 & 9) • Related to the previous one. • Getting into social behaviors • Interfacing with others of the same species. • Recognition, cooperation, etc. • We have to “deal” with relations!! Next? • 4) Foraging!! Looking for the energy that keeps us alive!! (chap 10). • 5) Closely related is anti-predatory and predatory behavior (Chap 11). Learning how to avoid being eaten can come in handy! Still more • 6) Aggression: (Chap 14). Learning how to be and when to be aggressive has survival advantages! • 7) Fear! Not listed but is a very helpful behavior to learn, especially in anti-predator context! • 8) Play (chap 15) what is its role? So there • As mentioned, these cover in a broad sense all that an animal learns and as we will see all play a role in aiding in the survival of the individual and the species. • Before we go to these behaviors, need to cover that other form of learning: Cultural transmission Cultural transmission (chap 5) • Talked about how an individual learns from external stimuli • What good is learning something new if you can’t share it!! • Why share it? Can aid in survival of relatives!! So sharing info has a survival advantage for your genes!! Cultural transmission • This sharing of information is cultural transmission. • Defined: Transfer of information from one individual to another via social learning or teaching (can be within or across generations) • As mentioned: this is a form of evolution without genes! Cultural transmission • Form of learning: social learning • Obviously similar to individual learning in terms of pathways, etc. • But involves learning from others • Lots of non-human examples: • Japanese macaques most noted • First was when one female learned how to wash her potatoes to remove sand and dirt Cultural transmission • Others followed suit and began to wash their potatoes. • Since then many more incidences noted. • • • • Characteristics of cultural transmission: 1) can spread quickly 2) more likely to spread to younger ones 3) seems to be some resistance to adapt in older individuals!! here Importance of cultural transmission • How does it differ from individual learning? • Individual is often trial and error • In cultural transmission, emphasis is: individual to individual • Way of transmitting individual learned behaviors, even after initial learner has died • Way of building on experiences: one learns and transmits a behavior, others improve it Social learning • As mentioned, basis of cultural transmission is social learning. • Various forms of social learning, two common ones are Imitation and copying Imitation • Must involve new behavior learned from others and must involve new spatial manipulation: Macaque and potatoes. • The keys are a new behavior and then undertaken by learner elsewhere Copying • One animal repeats what it sees the other do. • Differs from imitation in that need not be a new behavior. • Subtle difference but… Teaching in animals • So far, talking about an animal being observed doing something and then it is imitated or copied by others • No intent on the first animal (the model) in transmitting this information, passive role • Evidence that many animals take active role in transmitting information: Teaching • Implies one acting as instructor and others acting as students. Teaching in animals • Teacher must provide immediate benefit to students but not to itself. • Must “teach” only to naïve individuals • Must impart new information faster that students might otherwise receive it. Examples of teaching • Female cat bringing live prey to young • “helper” meerkats removing stingers from scorpions and then letting young catch and eat them • Many more usually females “teaching” their young. • This is a common theme found Other commonalities? • Usually are either “opportunity teaching” or “coaching” • Opportunity teaching: teacher places student in situation conducive to learning • Coaching: Teacher directly alters behavior of students by incentives. What next? • Modes of cultural transmission (direction of the transmission) • 1) Vertical • 2) Horizontal • 3) oblique Vertical cultural transmission • Information is transmitted across generations from parent to offspring. • Examples: Lots! All that “kids” learn from parents • Male birds learning song from good old dad • Females learning song preferences from mom! Horizontal cultural transmission • Learning from your peer group: Many animals segregate based on age (like humans!) and information is passed among them. Oblique transmission • • • • Across generations but not parent/offspring Young get info from nonparent adults. Common in species with little parental care Many of our fears seem to be acquired this Summary • Cultural transmission is a way of passing information on to other individuals • Avoids reinventing the wheel • Way of improving the wheel! • Can be done passively: social learning • Can be done actively: teaching (opportunity/coaching) • Occurs along different pathways: Vertical/horizontal/oblique Next: communication behavior (Chap 12) • Why we jumping ahead? • Saw that a lot is learned via cultural transmission • This transmission includes sight AND sound! • So, logically next behavior to consider would be communication! • Communication includes sight and sound Communication behavior • Defined: transfer of information from a signaler to a receiver. • Many of the communicative behaviors are part of the larger behavior we will be looking at: stress calls are part of “anti-predator” behavior • Will not concentrate on the overall behavior (which we will cover more in detail alter) • But the role communication plays in it. What is important? • Is it honest?? • Are the individuals accurately conveying information or are they trying to manipulate others? • Not just a problem with humans!! Honesty • Answer is yes and no. • Sometimes communication is “honest”, conveying correct info • Sometimes use communication to “get what you want”! When to be honest? • Classic ethological approach: both parties will be benefiting from information exchange • With this, less exaggerated signals (conspirational whispers) • Should be used in kinship communication • Unrelated, often hard to envision when both would benefit. • Should occur when impossible to fake information: Toads, only large males can make deep croaks • When costly to fake: length of song, energy intensive and only healthy ones can do it. When to lie! • Dawkins and Krebs suggested: most times!! • Communication in general is to convince recipient of something. • Advantage to “beefing up” our resume! • Selective advantage to convincing recipient you are something more than what you are. • Typified by exaggerated signals Recipient?? • Puts pressure on recipient! • Selection for those who can see through the lie! • Mind readers and arms race • The better recipient gets at detecting a fake, the better the fake has to get! Summary on honesty • Communication behavior is to convey info between signaler and receiver • Honesty may not always be the best policy • If send dishonest information, initiates an arms race • Depends on what trying to communicate • AND cost/benefit to honesty or dishonesty. Reasons for communication • Book: effort to solve problems that animals encounter in natural environment • Help see costs/benefits to communication • Three main problem areas: • 1) foraging • 2) mating • 3) predation Foraging communication • Problem: how to coordinate group foraging? • First… Why? • Social species easy to see: work as group and can more efficiently use a food source • • • • • Not so social species? Cost/benefit Cost: you have to share food! Benefit: if you are “recruited” to food source. Benefit: Recruiter? Keep track of food source if you have to leave and food source moves. here How coordinate? • Special food calls: many species of birds signal food source by calling • Crows: • Swallows • Physical movements • Waggle dance in bees • • • • • Chemical/vibrational communication: ants Recruitment phermones: Methyl 4-methylpyrrole-2-carboxylate 3-ethyl-2,5 methylpyrazine Placed along trails • Vibrational communication • Stridulation: Moving the ultimate body section (Gaster) up and down. • Found this conveys info to others about leaf quality. Problem: how to find a mate? • As mentioned 2 chapters on this but here we look at it relative to communication behavior • Two main communication pathways are: • 1) vocal • 2) tactile Vocal communication and mating: bird songs • Many species use vocal communication to find and attract a mate • But birds seem to have brought it to a well defined level: frogs croak, elk bugle, but birds sing! • Some simple but many quite complex • So will look at them as an example Birdsongs • Used for many things species can have one, 10, 100, a 1000 different songs. • But here we concentrate on those used for mate selection • Question: How songs are used to attract a mate. • Question: Why does it work?? Genetic filter • Thought that songs are a genetic filter females use to select mates. • One filter may be the number of songs a male can sing! The more songs, the more successful he is! • Why? More songs: older male or more fit to sing them! Harder to fake! • Remember, in communication receiver is trying to detect fake! Song complexity • Another filter is song complexity: • Evidence suggests that it is not what you say but how you say it! • Frequency, amplitude, rate of pauses, maximum note length, etc. All important components that transmit information about mate suitability • More energetically costly, more likely song is honest. Tactile communication and mating • Water striders: produce ripples in water with their legs. • Like songs, produce different ripple frequencies and amplitudes for different communication. • Calling mates; courtship: sex discrimination, etc. Ripple communication • Each is different, travel across the water and interpreted by females, which then respond accordingly! • How they use these to determine whether sender is honest or not, not clear yet but likely similar to sound waves of songs Problem: warn others about predators • When feeding in groups is beneficial, would be nice to warn others of impending danger! • Problem is: how, when, and why • How: mostly vocal calls (alarm calls) • Have the advantage of being very specific and can be transmitted rapidly When do you warn others? • Studies of birds show not always!! • Often only intersexual: mates will warn partner but not others of same sex! • Mate investment • Put a value on who to warn! • Indicates the potential “Why” • - when there is potential advantage to you or your genes! Honesty and alarm calls • We would assume that alarm calls would be “honest” • Most cases they are, transmitting important information • However, in some species we find alarm calls being used in a deceptive manner • The boy who called wolf!! Deceptive alarm calls • When would it be advantageous to use a deceptive alarm call? • Vervet monkeys: low ranking males will do so in potentially dangerous intergroup encounters: to avoid a fight! • Male swallows will do so if they see their mate with another male!! Does it work? • Initially yes! • But like the little boy, if you do it too often, others will ignore you. • Ground squirrels: If not backed up by seeing predator, will ignore calls How many false alarms should it take! • Obviously there is a danger in ignoring an alarm call: IF the villagers would have listened that last time…. • But how many false alarms are needed? • Depends on danger level, kind of like mimicry: • Depends on sender! Others may get to know one is unreliable. Summary communication behavior • Important in transmitting information to others. • Can be honest: what you say is what is so. • Can be deceptive: trying to manipulate • Is used to solve problems individuals face. • Depending on problem, depends on degree of honesty. • If dishonest, can start evolutionary race! Next: sexual selection (Chap 6) • Ok, have basic behaviors covered, innate, learning behavior, communication, etc. • Ready to look at “specialized” behaviors • Ones designed to deal with specific aspects of an individual’s life. • Again, remember will be a lot of overlap and at times may seem repetitious. • Can’t pigeon-hole behaviors Sexual selection • First we will deal with is behaviors aimed at acquiring more energy for your species: reproduction. • Various aspects of it, first is selecting who you will do it with!! • Believe it or not: don’t want to do it with just anyone! Gene selection • Reproduction is for passing on your genes • Evolutionary advantage to do it wisely! • • • • When we talk about sexual selection Consider: Intrasexual selection: same sex competing Intersexual selection: selecting mate Intrasexual selection • Darwin: competition for mates important part of reproductive success! • For most species: males do the competing, at least we think so!! • Why? Male competition • Why: • Males produce millions of sperm and can potentially inseminate many females at a low energy cost. • Females: produce 1-<15 eggs/cycle and although can and do mate with more than one male, really only need one male to fertilize eggs. • Also, energy intensive post fertilization here Bateman’s principle • 1) females should be choosier be eggs expensive to produce and reproductive success limited compared to male • 2) should be greater variance in reproductive success of males. Because there is an evolutionary advantage to being a successful male, evolution would favor competing for success. How do they compete? • Many forms: familiar with many • Direct fighting is common Male Competition • Displays: birds good at this! Mate choice • However, not all just male competition. • What is the female’s role? • Used to think it was passive: waiting patiently on the sideline to see who the winner would be! • But females have stake in this! • More so than males: it is their ONLY chance at passing their genes on! So who is choosing whom? • Answer: both really. • So we now talk more about Mate choice • we look more not just at the glamorous battles/displays of the males but more at the often subtle reasons WHY males do what they do, which is often driven by female choice! • Colorful male birds! Male secondary sex characteristics • Why do male lions have a mane? • Why do male baboons have such red butts!! • It is because it is what it takes to convince a female! So who is doing the choosing?? • Mate choice is a male trying to convince female he is the right one and females evaluating IF that is really the case! • So we need to look at evolutionary models of mate choice to better see how and why we males have gotten so ridiculous!! Evolution of mate choice • Past: looked at flashy males • Today: lots of research into female choice • Underlying assumption of most earlier work is that female choice is under some genetic control – there is genetic variation in female mate choice behavior. • Four evolutionary models proposed: Evolutionary models Direct Benefits model • Natural selection favors females who are genetically predisposed to choose mates that provide benefits beyond just good genes. • Female will choose mate that provide important resources: food, safe shelter, assistance, etc. Evidence for this model • Female scorpionflies: choose mates based on the size of prey item they bring. The bigger the better, if you don’t bring anything, you get rejected! • Nuptial gifts: she eats while they copulate. More • Female determines how long they copulate based on size of nuptial gift: again, larger longer • Longer they copulate, more sperm transferred • What does she get from all this? • Larger gift, longer copulation time, more eggs produced. • Also provides a meal for her! Genetic link • Strong selection pressure on males who can bring large nuptial gifts. • Finding large gifts is time consuming and dangerous. • Few can do it, so they do the most mating • Some steal prey by mimicking females • Evidence that females select males based on resources provided and how it can affect male foraging behavior. Good genes hypothesis • Concentrates not on resources females get from males but the quality of the sperm they receive: good genes! • Good genes: those that code for some suite of favorable traits, which are passed on by a choosey female. • More indirect benefits than previous • Increased survival of offspring But…how do you tell? • May sound good and field evidence that females selecting for other than direct benefits. • E.g. Pronghorn, females select male based on size of harem…larger he is able to keep, more “attractive” he is. • But how do females know this is good genetically?? Basic questions • 1) How can females determine which males have “good” genes? • 2) If she is using some criteria, how does she know the male is not faking this criteria?? That communication thing again! Honest males, do they exist? • • • • Answer the second question first. Saw in communication behavior Selective advantage to a male being dishonest But…selective advantage to female who can tell the difference! • Arms race that should lead to Honest indicators of gene quality…but what are they? Honest indicators of gene fitness • In general two major factors hard to lie about. • 1) survival capabilities • 2) physical health • Can be related and both are energy intensive. Survival capabilities • How do you tell that your potential mate can give you genes that would enhance survival of offspring? • You test them!! • Not surprising that for many species, males have elaborate courting rituals. • Many related to health as we will see. • But some are tests of survival. Survival tests • Usually say brightly colored male birds are so to “attract” a mate. • Why attractive? Assume is a good indicator of health. • But, could be a test against predation!! Survival test • Elaborate courtship rituals: • Brightly colored, singing, and dancing like a fool!!! • If you can survive predation attempts, you may have what it takes! Survival • Frogs, fireflies, list goes on and on of males putting themselves in danger to attract a mate. Why? • You would think that evolution would favor safer courtship rituals unless its demanded by the females! State of Health • Majority of courting rituals (usually male-male competition) can be considered ways females can judge health of potential mate. • Not surprising that many involve physical contests. • Definite filter regarding health of male and hard to fake. Physical contests • Leks in grouse, contest for preferred central areas: physical strength. • AND safety for females for mating! State of Health • Brightly coloring • Takes energy to produce AND maintain all this. • What is the first thing to go when you feel sick? Personal hygiene! • So two tests: • 1) do you have genes to grow them/ 2) Are you healthy enough to maintain them? Bright colors • Redwing black birds: Bright colors • Bright colors may indicate parasite resistance; Sticklebacks Courtship rituals • Besides putting yourself in danger, it takes energy to dance around, hop up and down, sing, build things (bower birds). Odors! • Evidence some females use odors as indicators of health – good genes. • MHC: Major Histocompatibility complex • What? • Set of genes related to disease resistance • These are “good” genes to get!! • Combination of genes in this complex result in different body odors! MHC • Found males with dissimilar MHC combinations were more attractive to females • Also, different odors related to number of genes in this complex and females attracted to males with higher numbers of genes! here Symmetry • Symmetry of body form: similarity of left and right sides. • Related to developmental stability • Which is related to ability to adapt to change. • Obviously equating health with genetic well being.