Chapter 13: WHAT IS COMMUNICATION? - When one organism transmits a signal that another organism is capable of responding to appropriately. • Information is transferred from a sender/signaler to a receiver • SENDER → *SIGNAL* → RECEIVER • The ‘SIGNAL’ provides information. • BOTH THE SENDER AND RECEIVER MUST BENEFIT COMMUNICATION IN TERMS OF EVOLUTION - Favours “SENDERS” very specifically to increase their own FITNESS WHAT IS A *****SIGNAL? 1. ‘behavioral, physiological, or morphological characteristics fashioned or maintained by natural selection because they convey information to other organisms’ 2. ‘act or structure which alters the behaviour of other organisms, which evolved because of that effect, and which is effective because the receiver’s response has also evolved’ (modern definition) ANIMAL COMMUNICATION “*******WHAT information do animals communicate? • Identity (individuals) - Just like we have a name, other species also have their own identities (song birds- specific contact calls) • Mood • Intentions (e.g., fighting) • ENVIRONMENTAL factors: - Location of food Potential mates (MATING) PREDATOR THREAT TYPES of communication: • Visual • Auditory • Chemical - (ANT PHEROMONES VIDEO) squeezed out the pheromones out of one ant and laid out a trail of his own- other ants followed and piled up on the trail • Electrical • Tactile/Thermal • Vibrations “MATING BEHAVIOR OF A POND SKATER / WATER STRIDER (GERRIDAE)” - VIBRATION EXAMPLE THROUGH SURFACE WAVES MATING BEHAVIOUR - Experimenters were able to use a dead male and tapping it specific body part, creating the vibrations on the surface of the water… ATTARCTING A FEMALE ACTIVE VS. PASSIVE COMMUNICATION • **ACTIVE process - EFFORTFUL PROCESS - e.g., whale echolocation e.g., deer antler fights (smashing antlers together) teacher teaching • **PASSIVE process Colouring - e.g., poison dart frog (predator see the vibrant skin and should recognize if it ingests it, they will become ill or die) e.g., bees & wasps e.g Scarlet snake looks venomous - moth looks like Bee/wasp Plumage - Dull plumage = signal for poor quality (WILL NOT FIND MATE) e.g., peacock’s bright colours - if they don’t have them, they will not find a mate) WATER STRIDER EXAMPLE OF WHAT WE LEARNED JUST NOW: INTRASPECIES COMMUNICATION (human to human) (dog to dog) (cat, to cat) INTERSPECIES COMMUNICATION - exists BETWEEN SPECIES • PREY-TO-PREDATOR interactions - “Don’t try to eat me because… I am dangerous” orrr: - Skunk posture Colouration/coloration = poisonous Armor = protection or painful (shells, stingers, quills) •PREDATOR-TO-PREY interactions - Changing the behaviour of the prey to make them easier to catch (E.g. ANGLER FISH in finding nemo) 1. HOW DOES COMMUNICATION EVOLVE? • Evolution would favour senders that signal in order to increase their own fitness. - At the expense of the receiver! Communication then would be defined as an “attempt by the sender (“manipulator”) to manipulate the recipient in some way.” • Evolution would also favour RECIPIENTS who could detect when they were being manipulated (“mind readers”). - And only act on information that benefited them! • Thus, we have a “COMMUNICATION ARM’S RACE” with ever-better manipulators/mind readers! • Krebs and Dawkins (two famous ethologists) argue that both types of communication exist—a ‘classic’ cooperative signal exchange and the manipulative type: - The MANIPULATIVE/MIND READER COMMUNICATION TYPE may use conspicuous, exaggerated signals (think: ‘used car salesman’). The COOPERATIVE TYPE may use quieter, less obvious signals (called “conspiratorial whispers”) • DECEIT is a special form of communication. (DO ANIMALS LIE?) - - This is common across species (e.g., predator/prey communication), but not within a species. e.g., mother kildeer bird pretending to predator to have a broken wing • Lying is only an advantage when everyone else is honest! - Also, selection favors the ability to detect deceit. Signaller will benefit from lying, receivers will benefit when they are detecting the dishonesty HANDICAP PRINCIPLE: has to do with using a lot of resources to be deceiving. E.g. (energy) injured gazelle - jumping up and down pretending it is healthy - DECEPTIVE USE OF ALARM CALLS BY MALE SWALLOWS, HIRUNDO RUSTICA: A NEW PATERNITY GUARD • male swallows faking alarm calls to increase the certainty of paternity. • Deception can increase certainty of paternity using paternity guards! • Maximize reproductive success by caring for their own offspring and attempting to fertilize the eggs of additional females without providing any paternal care • Males were able to disrupt extrapair copulation attempts toward their mates by giving alarm calls. - Causing all nearby swallows to take flight! • Solitarily breeding males gave alarm calls when their mates were absent from the nest if a male mount was previously near nest during and before the egg-laying period. 2. HOW DO SIGNALS CONVEY INFORMATION? • Communication is completed through the exchange of signals! • A signal is the physical form in which a message is coded for transmission through the environment - DISCRETE SIGNALS: On or Off (e.g., fireflies) GRADED SIGNALS: Vary in intensity (vary from low, medium, high pitch) • Signals vary in modality: - **VISUAL SIGNALS work over shorter distances, but can last for a long time (e.g., coloration, antlers for mate choice). (e.g mallard colouration - males are different) **AUDITORY SIGNALS can be short or long-range (e.g., long-range elephant vocalizations vs. short-rang gorilla grunts). **CHEMICAL SIGNALS can last a really long time (e.g., pheromones or scent trails - urine with big cats). Signals can be quite complex: - Two or more signals can form a COMPOSITE SIGNAL- more than one building up (e.g., tigers with ear position and vocalization) https://www.youtube.com/watch?v=LL99pufzHjo&list=PLbPhJdEXzeOg99UlZHbLalQN 9omvQr0o8&index=3 (TIGER VIDEO) Tigers communicate with smell, hearing, sight. TIGERS RUB THEIR CHEEKS ON EACHOTHER (rubbing scent glands) they also have the urine scent TIGER CHUFS/CHUFFING IS A GREETING TIGER SPOTS ON BACK OF EAR - if they rotate the ears and u see the spots when infront of the tiger.. it is very bad for you.. you will be eaten alive • Context-dependent - same signal in different situations means different things! - Chickadee use the chika deedeedee for various reasons ( food or predators) the more deedeedees mean there is a predator • Some species use ‘META-COMMUNICATION’ to signal the context of behaviours that will follow (e.g., play bowing in dogs or plaay face with gorillas). - https://www.youtube.com/watch?v=rlSTwNw4Y9U&list=PLbPhJdEXzeOg99UlZHbLalQN9omvQr 0o8&index=6 3. WHAT ARE THE FUNCTIONS OF COMMUNICATION? • The ultimate function of any behavior is to increase fitness (i.e., reproductive success)! • But communication can increase fitness in a variety of ways: 1. Coordination of feeding: • e.g., Group-spacing and coordination in wolves - Howling to coordinate everyone coming back together (acoustic).. Visual cues as well as rubbing each other • e.g., Crows and ravens “yelling” when they find food • e.g., Rooster alerting ‘his’ hens about finding food 2. REPRODUCTION and securing a mate: https://www.youtube.com/watch?v=gqsMTZQ-pmE&list=PLbPhJdEXzeOg99UlZHbLalQN9omvQr0o8&index=8 • The best examples of this involve the evolution of song in birds! • Birds do NOT have a larynx (like humans do!), but rather a syrinx that produces sounds. - The syrinx has two compartments (left vs. right) and the brain controls each independently! e.g., Lyre bird of Australia https://www.youtube.com/watch?v=VjE0Kdfos4Y (VOCAL MIMICRY) 3. Social status and aggressiveness - https://www.youtube.com/watch?v=YvP3aHiDDn8 - ^flat lizards showing their social status with stomach colour 4. Detecting predators • Alarm calls are to warn conspecifics about the presence of predators (e.g., Vervet monkeys). • Alarm calls can also be used as deceptive communication. VERVET MONKEYS: 3 distinct alarm calls below • Eagle alarm: Chuckle then dive into nearby bush • Leopard alarm: Loud bark then run up nearby tree • Snake alarm: High-pitched chuttering, standing up on their hindlegs and mob the snake. 5. Parent/offspring communication (e.g., begging, synchronization of hatching, etc.) - Baby birds are known to vocally communicate in their eggs with other unhatched babies to let them know they are coming out. 4. CHANNELS OF COMMUNICATION 1. Odor: • Used by almost all animals, except for birds! • Pros: Long range, works at night, long-lasting, low cost • Cons: Slow transmission rate, difficult to identify sender, needs to be within certain parameters • Pheromones: Secreted or excreted chemical that triggers a social response in members of the same species (SPECIES SPECIFIC) - SIGNALING PHEROMONES: Provides information about the individual RECRUITMENT PHEROMONES: Attract and orient activity (usually to a trail - ANTS) • Pheromones often produced by glands or in urine; response by receivers is often ***FLEHMEN • e.g., Horses, giraffes, felines… - https://www.youtube.com/watch?v=SY1hbyHim6s&list=PLbPhJdEXzeOg99UlZHbLalQN9omvQr0 o8&index=10 horse makes a face reacting to the scent (FLEHMEN) https://www.youtube.com/watch?v=0YezFh4RMY&list=PLbPhJdEXzeOg99UlZHbLalQN9omvQr0o8&index=11 tigers flips top of mouth to snort/squish the scent (FLEHMEN) - special organ top of their mouth that does this 2. Sound • Used by almost all animals • Pros: Can be long range, lots of variety, works at night, transmits quickly, can identify sender • Cons: Fades quickly, high cost (bc it tells predators where you are/ also takes a lot of development over time) • Acoustic communication varies, including low frequency sounds (e.g., kangaroo rats, elephants)- to ultra-high frequency sounds (e.g., canids, bats, rodents) • Sound can also be used underwater (e.g., marine mammals)! • https://www.youtube.com/watch?v=JODHKHzr85U - Dolphins communicate through blowhole (whistles and clicks) SONGBIRD VOCALIZATIONS: these are opposite of chickadees • Calls - “Brief, monosyllabic bursts of sound” Individual recognition, courtship, motivational states, predator information • Songs - “Repetition of a sequence of syllables” Males defending territory or attracting females CHICKADEES: VOCAL LEARNERS (learn to survive through vocalizations from a model/parent) A type of North American songbird • One of only six groups of animals that are considered ‘vocal learners’ • Humans are vocal learners - Thus, chickadees provide a strong comparative model for language and cognition! • Produce several vocalizations critical to their survival: • Chick-a-dee call • Fee-bee song (AKA CHEESBURGER, or HEY SWEETIE) • Gargle call (ANTAGONISTIC) UNHAPPY BIRD 3. Tactile (i.e., touch) • Used by almost all animals for at least some types of communication • Pros: Works at night, low cost, can identify sender • Cons: Short range only, fades quickly • In mammals, touch is used during copulation (e.g., lordosis in rodents, stimulation of ovulation in species like cats, and grooming in primates). • Touch is often used by invertebrates through antennae (e.g., cockroaches or bees). (HONEY-BEES - Communication of food location Round dance vs. Waggle dance) - round dance: food 50 meters away circular movement - waggle dance: vertical of hive. Do circular figure eight patterns. Direction is indicated towards that vertical (food 100 meters away) 5. Visual - Used by almost all animals (NEED FUNCTIONAL EYES THOUGH) • Pros: Medium range, lots of variety, does not often work at night (except fireflies!), transmits quickly, can identify sender • Cons: Fades quickly, medium cost • Visual communication includes a ‘visual display’- long sequence of a bunch of different behaviours • COMMUNICATION BEHAVIOURS (e.g., visual displays) probably evolved from noncommunicative behaviours, like INTENTION MOVEMENTS (indication about subsequent action) or DISPLACEMENT ACTIVITY (occur in conflict situations) E.g. 2 cats looking like they about to fight, then nothing happens and then they lay down too much arousal occurring causes behaviour to stop drastically (Switching behaviour when there is too much arousal/conflict) LANGUAGE VS. COMMUNICATION • To what extent does animal communication resemble language? Language is potentially the most important intellectual capacity possessed by humans. • Live in large social groups • Teach skills and information • Express our feelings and thoughts ****************WHAT IS LANGUAGE?**************** ***Arbitrariness of - Discrete units (e.g., words) A single object can be referred to by many different words/languages ***Semanticity - (vervet monkeys, chickadees, songbirds arbitrariness and semanticity) Specific meaning ***Displacement - units - difficult to find in other nonhuman species (HONEY BEES - flower patches) Communicating about events in another time or space (talking about what you did today at dinner) ***Productivity - (song birds) Grammar/syntax – rules of production Create many sentences with a limited vocabulary LANGUAGE ABILITIES • Attempts to teach orangutans, chimps, and gorillas to speak English - Failure due to vocal tract constraints! (structured differently than us) Layrnx is different in apes, cant physically speak words ARTIFICIAL LANGUAGE • American Sign Language (Ameslan) - Cheremes (like phonemes) – one gesture Washoe (180), Nim (125) & Loulis WASHOE TAUGHT LOULIS SIGN LANGUAGE WITHOUT THE KEEPERS • Plastic Tokens - Plastic objects varying in shape, size, colour, and texture (arbitrary but have semanticity) • Lexigrams - Symbols serving as words Seen with bonobo • Spoken English (requests) - Kanzi – “Take the potato outdoors” (72% accuracy) CHAPTER 14 HABITAT SELECTION, TERRITORIALITY, AND MIGRATION Habitat: - Answers a WHERE question WHERE can I find this animal and WHY does it choose to be there? e.g., Distribution Patterns Territoriality: - Answers the HOW question HOW does this animal succeed in this landscape? Exclusive access to food, mates, etc. Migration: - Answers a WHY question WHY move away from the tropics for the temperate summers? Funfact: sometimes birds go from hot summery places and go to cold places up north HABITAT SELECTION Habitat can affect the behaviour of the individual. - e.g., anti-predation defense, foraging strategy, mating choices, etc. Animal behaviour can affect the choice of habitat. - For mobile animals, there is some choice of where to live… What determines the attractiveness of one vs. a second habitat “patch”? HABITAT SELECTION FACTORS: What affects the choice of where to live? ABIOTIC FACTORS - e.g., shelter, temperature (especially for ectotherms-dependant on weather ), water, wind, etc. (NOT LIVING) - Would want to live near water Wind can actlly blow ur scent towards predtors BIOTIC FACTORS - e.g., food availability, predators, mating opportunities, parasites, etc. (LIVING) IDEAL FREE DISTRIBUTION - predicts where animals should be found, based on available resources. The red line on the graph is the point called the “Nash Equilibrium” where resources are equal between patches. LOTS OF STUDIES • The ideal free distribution has been extensively studied in field and lab experiments. - e.g., stickleback fish experiment - they go where there is food, unless there is too many fish, bc of this, some move onto to new patch Basically, the number of individuals in a patch should match the number of resources available in that patch! ANOTHER BIOTIC FACTOR IN HABITAT SELECTION: PARASITE AVOIDANCE Natural selection should act strongly for behaviour that reduces exposure to disease. - Avoid pathogen-heavy habitats (e.g. ticks - animals learn about them and avoiid them) Avoid sick or infected individuals (e.g. red chested bird video we watched - tuberculosis) Example - Study of ovipositing gray treefrogs - Frogs selected ponds with no snails or uninfected snails! BRAINS ARE IMPORTANT IN HABITAT SELECTION! SPATIAL MEMORY (memory for location) can be an essential ability for animals to select habitats. - e.g., Remember where food is cached, where their dens are, where predators are, how to navigate through their environment Note: If you’re interested in some of these cognitive abilities, take seminar course in Comparative Cognition (PSYC 4285)! ***TERRITORIALITY Once you selected a habitat, you must achieve territoriality! Territory refers to an area that is DEFENDED vs. a place where an animal simply lives, which we call a “home range”. **HOME RANGE (NOT DEFENDED): Area that is visited with regularity but not defended (this is very common in animals). - Animals of the same species might share a home range or have overlapping home ranges. They often avoid contact with other groups although sometimes they might mingle (e.g., breeding or migration; i.e., groups of elephants). Some species don’t have a home range or a territory and are simply ‘free ranging’ (e.g., army ants)! HOME RANGES Vary in size depending on whether the animal is living in urban or rural areas! - e.g., Raccoons in the city have home ranges of about 3 square blocks, while ones in the country can have ranges of 20 km! Also, can vary between the sexes! - e.g., Males often have larger home ranges, and overlap the home ranges of several females. HOME RANGES IN RACCOONS GPS collars on five 5 raccoons in downtown Toronto. Collars recorded the location of each animal every 20 minutes so lots and lots of data Where did they go? Preferred backyards / private houses over busy main streets **TERRITORY: Area occupied and defended against intruders and conspecifics. - For territorial species, the defendable territory will be a smaller subset of the larger home range! Various kinds of territory: - **ALL PURPOSE: Where all of the life history of the individual can happen **SPECIALIZED: Mating (e.g., lekking), nesting, roosting, wintering, etc… COST-BENEFIT OF HOLDING A TERRITORY: - Benefits should be higher than cost Bigger territory size has bigger value and could come with lots of costs (like energy, more to defend - predtrs) Lewa Wildlife Conservancy Grevey’s Zebra - highly endangered - Lions are their preddators Looks at the home rang of lions and zebras TERRITORIALITY AND LEARNING Experiments with lizards (Stamps, 2001) - Examined juvenile lizards and decision mechanisms for determining territory What was important? - Safety from predators & optimal temperature How did lizards decide? - CONSPECIFIC CUING: The choices of others! they watch others and learn from others. **MIGRATION AND NAVIGATION Migration happens in all animal taxa! - i.e., Birds, mammals, insects, fish, crustaceans, etc.. Different levels of migration (i.e., Migration Continuum) Examples birds: - OBLIGATORY: mandatory - they absolutely have to do it (e.g., insectivores - warblers( or hummingbirds for nectar (sea turtle - obligatory migration bc they have to mate/have offfspring) FACULATIVE: u don’t have to, u can if u want (e.g., robins, cardinals) IRRUPTIVE: if something shows up all of a sudden, they will migrrate e.g., crossbills or snowy owls) due to lack of food (e.g., pine nut-dependent, rodent-dependent) *************REASONS FOR MIGRATION************* FOLLOWING FOOD SUPPLIES - e.g., Wildebeests follow the rains to find phosphorus rich grasses e.g., Passerine birds nest in the Boreal forest in a superabundance of food (e.g., mosquitoes) AVOIDING WEATHER - e.g., Caribou migrate South to avoid the snows and the lack of food PROCREATION: Mating, nesting, rearing, etc. - e.g., Whales will have calving areas that are safe from predators ELEPHANT MIGRATION Save the Elephants: Elephant tracking https://vimeo.com/48959100 - Strict migrational patterns, have a route they take every year traditionally Farmers creating farmland (or roads) effect their routes When they raid the farms, they get killed sometimes bc they can eat an outstanding amount of crops Mfuwe Lodge: Looking for mango https://www.youtube.com/watch?v=o2IQ2A7Orc8 - Running through hotel to get mangos MALE ELEPHANTS are MORE AGGRESSIVE ELEPHANT CORRIDORS - highway under pass allows them to safely cross roads - Get elephants to go through corridors with either elephant dung or vanilla scent (they love it) STUDYING BIRD MIGRATION Point Counts (see image) - Individuals sit and count birds Monitoring surveys RAPTOR MIGRRATION: a lot of TIMING - Each species have different variables for migration Bar-tailed godwit migrrration - Over 10,000km non-stop migration every season. It takes 9 days! *************MIGRATION TRIGGERS IN BIRDS************** Length of day (i.e., PHOTOPERIOD) - longer sunny days - Linked to hormones (experimentally castrated males act as controls!), including the pituitary gland - (if we remove this they will act different) TWO-STAGE: Prep then go! - 1. Increased fat deposit (prep stage) 2. “ZUGUNRUHE” (Migratory Restlessness), often at night, evidence of a powerful internal clock MIGRATORY RESTLESSNESS (“ZUGUNRUHE”) (INNATE) - Some aspects of migration are heritable (i.e., genetic). Onset of migratory activity between laboratory vs. wild birds (similar!) - Captured wild birds and selected for “late-onset” migration restlessness via SELECTIVE BREEDING. Within two generations, the offspring had shifted their “zugunruhe” by more than a week (via GENETIC HERITABILITY) BUTTERFLY MIGRRATION “Monarch migrrration” https://vimeo.com/41075641 - Inherited multi-generational routes (10,000 km) Using sun compass (time-shifting experiments to test solar navigation), magnetic fields, biological clock, etc.? Researchers are still trying to understand how they do it! DIAPAUSE: Fat, protein, and carb storing Darker colour & bigger wings (compared to summer monarchs) MIGRATION: COST VS. BENEFITS Costs? - Well, fewer than 50% make it back!! Energy requirements (40% drop in weight pre/post) - Weather and other obstacles Predators and parasites (twice the variety!) - Migrants invest more heavily in immune function (Moller & Erritzoe, 1998) Competition for food in wintering range Competition for territory every spring EVOLUTION OF MIGRATORY BEHAVIOUR Evolution: How did migratory behaviour likely evolve? There must be BENEFITS!! - Reproductive advantage (i.e., super abundant food supply) Reproductive synchrony/all together (i.e., overwhelming predators bc there are so many of u) ALTERNATIVES TO MIGRATION Winter foraging - Possible for seed & fruit-eaters Not possible for insectivores! HIBERNATION: Bear den & Bear hibernation - Pineal gland allows us to integrate daylight changes. . BEARS HAVE A VERY SMALL PINEAL GLAND - Raccoons do not hibernate *****ORIENTATION - Words to remember: KINESES (general movement) & TAXES (specific directed movement) Negative phototaxis example: NEMATODES (moving away from light) Positive phototaxis example: MOTH (moving towards light) Other taxes: - Gravitaxis (spiders), (moving towards gravity) Chemotaxis (E.coli), (moving towards chemicals) Thermotaxis (slime molds), etc... (moving towards heat) HOMING PIGEONS - u can take one and drive them far away, they will know where to fly to get back home (without knowledge or practice) - Simple retracing of the route Path integration - “if u went out in all these zigzags, u will want to just take a straight line home” Piloting (i.e., landmark use) Compass orientation: Using the sun as a bearing True Navigation: Need to know where they are and where they are going (two systems!) TESTING HOMING: - Displacing animals from home ranges Measuring “vanishing bearing” in homing pigeons Testing the importance of the senses by manipulation (e.g., blindfold experiments) ROUTE FOLLOWING Route following: Homing using chemicals cues (e.g., slugs, wood ants, chitons, caterpillars) - Chiton moving fast! Recognize landscape through primitive eyes? Magnetite teeth = magnetoreception? PILOTING WITH LANDMARKS: Fixed cues: - Construction of a “mental map” Tinbergen digger wasp experiment: Digger wasps https://www.youtube.com/watch?v=qm0Z0ceezQ&t=117s (2:33 - 4:39) mental map of fir cones / pinecones Migrating waterfowl follow coastlines Wood ants use skyline panorama Topographical piloting is NOT much help for migrants if: - Featureless landscape First time migrants Night migrants SUN NAVIGATION Frosted Lenses Experiment (pigeons cannot see very well with these lenses) - Pigeons were successful HOMING PIGEONS do NOT need to see landmarks as long as they can see the sun! Clock-shifting experiments manipulate daylight to fool subjects (Monarch butterflies) COMPASS ORIENTATION - Using the sun to maintain a bearing in an unfamiliar landscape! Ants (Cataglyphis) also use a stride counter to measure distance (i.e., “pedometer”) ^^(stilt or stumps experiment) they either cut off legs or added stilts, bc of this the ants migrration distance was doubled bc of longer legs There is a need for an internal clock to adjust for the sun’s movement in the sky (e.g., Monarch butterflies). Bats use the sunset position to orient themselves before their nocturnal migrration. CELESTIAL NAVIGATION Moon as a compass (e.g., moths) Stars used by night migrants (e.g., Passerine birds) - Planetarium experiments: manipulate which stars are visible (e.g., Indigo buntings). Young birds must be exposed to the night skies! Birds learn the star map not just the North Star! Night migrants orient using certain geometry of constellations and the North star polaris. OLFACTORY CUES - Manipulate odors! e.g., Salmon (fish) imprint on the odor of the natal stream. The role of olfaction in pigeon navigation is still debated, but some research suggests an olfactory map… Although anosmic(not being able to smell) pigeons make lousy homers!. - MAGNETITE in pigeon beaks and brain point to other systems at play. ^^Magnetic properties of earth MAGNETIC NAVIGATION MAGNETORECEPTION is often combined with other cues (Able & Able, 1990). - Experiments with pigeons with magnetic coils on their heads or bar magnets on their wings also confirmed magnetic navigation! MAGNETIC FIELD Sea turtles use a GEOMAGNETIC MAP to navigate the seas in combination with possible landmarks as they near the coast. Jenna’s ANT RESEARCH STUDY Navigation in a species of desert harvester ant - Vermessor pergandei - Sonoran Desert, Arizona Mirror experiment using reflection as “the sun”.. this fucked the ants’ direction and went opposite way SUMMARY OF STUDIES - Sun location matters a lot to them Pheromones also matter a lot to them Individual navigational abilities - Direction & distance of column (vectors) Backtracking dependent on celestial cues - Primarily the sun’s location Column pheromones act as contextual cues - Two-vector system NEURAL MECHANISMS Multiple navigation systems in use, and backups! Spatial learning is centered in the hippocampus (so is olfaction). LESIONS to the FOREBRAIN strongly impair the development of spatial abilities in young birds. Once learned, similar lesions do not reduce retention of mapping skills! Recent research describes neuronal responses in the pigeon’s brainstem that show how single cells encode: - Magnetic field direction Intensity Polarity SO HOW DO ANIMALS FIND THEIR WAY? Some species use trail marking to make their way home Others use path integration, keeping track of an internal egocentric map (self focused map) Long-distance migrants can use piloting with landmarks Many species use the sun or the stars as a compass to maintain a bearing Olfactory and geomagnetic cues appear to be playing a role in mapping Trials of Life: Finding the Way .. NOT MANDATORY http://www.infocobuild.com/books-and-films/nature/trials-of-life/episode-05.html chapter 15 AGGRESSION - studying this in animals can also explain aggression in humans Natural selection is based on a competition for resources: this is why aggre$$ion is rising. - e.g., mates, territories, food, etc. And competition sometimes results in aggression! Aggression in animals has been an important topic to philosophers for hundreds of years. Are all animals (including humans) innately aggressive, or are cooperation and altruism the norm? - ARE YOU BORN WITH AGGRE$$ON OR YOU LEARN IT? - Turns out both exist, depending on the situation (i.e., costs vs. benefits of fighting). DEFINING CONFLICT AGGRE$$ION VS AGNOSTIC BEHAVIOUR ******************AGGRESSION - Complex term, difficult to define. Intention to inflict noxious stimulation or destruction on other organism, or exclusion from resources (e.g., food, shelter, or mates) - very human term AGNOSTIC BEHAVIOUR - More specific term, referring to conflict AMONG CONSPECIFICS (ur own species) (e.g., territorial dispute, dominance, sexual, parental, parent-offspring; NOT ********COMPETITION predation) Basis of most agonistic behaviour is competition for a limited resource (e.g., space, food, water, mating opportunities, etc.). Two (2) kinds of comptition: 1. ****EXPLOITATION: Passively taking up the resource (e.g. hogging all the food bc others don’t have it) 2. ****INTERFERENCE: Active interaction to reduce another’s access to resources ( E.g. I am going to block you from getting the food/water) Generally, most species have evolved ways to reduce injury and death from conflict, so conflict behaviour is often seen in the form of threat, displays or contests. Rarely see “fights to the death” among conspecifics (although some conflicts result in injuries that lead to death, especially over access to mates). IN GROUP-LIVING SPECIES(shared resources, shared territory, better mate access) One way to reduce daily conflicts and to ensure relative calm in socially living species is to establish a dominance hierarchy, often seen in primates. - e.g., Rhesus macaques (a lot of behavioural observation methods come from studying these monkeys) ***DOMINANCE = One animal controls the behaviour of another animal - - If X is DOMINANT to Y, Y is SUBORDINATE to X. If X and Y compete over a resource, X will win/Y will forfeit: X > Y. FEMALE RHESUS MONKEYS - female dominant society - Dominance determined by mom’s rank All have specific ranks in the hierarchy Individuals get their mother’s rank.. a baby can be a higher ranking individual than an adult monkey MALE RHESUS MONKEYS - Dominance determined by seniority (how long they been around) LINEAR DOMINANCE HIERARCHY: Sometimes it’s more complicated: - Z and Y form a coalition, making them more dominant than X ^ Species specific Situational specific ****DOMINANCE There are costs associated with being dominant. - e.g., Need to fight and break up fights, defend resources, be constantly vigilant Why do it? - Benefits of dominance outweigh the costs! - Access to mating, access to food, etc. (black hat chickadee subordinate would fly away from feeder once the more dominnant bird lands.) IS NATURALLY SELECTED FOR May also have better immune systems https://www.youtube.com/watch?v=OYFRICCVK4M&list=PLbPhJdEXzeOgoXRi32mv1pMisPPKhNfsz&in dex=5 social rank and stress from changes in social rank alter GENE EXPRESSION IN PRIMATES (rhesus macaque) when social rank improved gene expression involved in immune function changed within weeks) RECONCILIATION Reconciliation: An increased probability of contact between individuals following agonistic interactions. These involve special behavioural patterns (e.g., reassuring, hugging, kissing; chimps). ON THE FLIP SIDE… Do animals ever purposefully kill each other? - Rarely! There have been instances of cannibalism, but this is seen in abnormal conditions like captivity or when food is scarce. Sometimes SIBLICIDE occurs, especially in hyenas. (eating younger smaller/siblings) parent off spring conflict Sometimes infanticide occurs under stressful conditions, or when new male moves into a group. - e.g., In lions, although this is controversial… To see some examples of conflict in a variety of species: Trials of Life: Fighting, https://www.dailymotion.com/video/x6vzaw3 not mandatory MECHANISMS UNDERLYING AGONISTIC BEHAVIOUR: why does aggression occur? 1. INTERNAL FACTORS Hormones - - Testosterone is an important one, but not the only one! You can measure this ^^^ with faeces / feces Neural mechanisms - Ventromedial nucleus of the hypothalamus, amygdala, serotonin levels (high levels in vertebrates = lowered aggression; although the opposite is seen in invertebrates) Genes - Controlling agonistic behaviours or hormone production/reception have been found in many species E.g. genes can be manipulated for more aggre$$ive individuals 2. External factors (environment - everything outside our body that’s happening) - PAIN and frustration can lead to aggre$$ive behaviour If you see an animal acting agonistic, checking to see if they are in pain is the first thing to check Social factors, based on prior experience (Learning to be more aggressive) PRIOR SOCIAL EXPERIENCE: 1. Winner and loser effects: if you win a fight, you will win more; losers of one fight tend to lose future ones (May have hormonal basis - testosterone backs them up and helps them keep winning) 2. Bystander effects: “Eavesdroppers” can learn about potential opponents 3. Audience effects: Having an audience changes the behaviour of the opponents (And SPECIFICALLY WHO is in the audience also changes their behaviour! e.g., chimpanzees) [Connected to SOCIAL FACILITATION ***] MOST ANIMAL CONFLICT DOES NOT RESULT IN FIGHTING! **Most conflicts are settled by THREATS, DISPLAYS, or CONTESTS. Why? - Game Theory models (Prisoner’s Dilemma/ TIT 4 TAT frm Cooperation lecture GAMETHEORY - explains the evolution and maintenance of aggression and why it occurs GAME THEORY MODELS - weighing the costs against the benefits All models of the evolution of fighting behaviour/aggression involve weighing the costs of aggression (e.g., time spent not doing other things, physical injury, even death) against the benefits of aggression (e.g., gaining access to food or a mate - valued resources). When is it worth it to fight, and when should you walk away? 1. The HAWK-DOVE GAME - This is the first and best known model of the evolution of aggression! Hawk = Aggressive strategy; escalates and keeps escalating until injured or obtains resource(s) Dove (also known as “mouse”) = Displays, but then backs down if opponent escalates Which strategy pays off depends on the value of the resource (V) and the cost of fighting (C)! - If V is much higher than C, hawks win; If C is much higher than V, doves win. The best strategy (Evolutionarily Stable Strategy, or ESS) in conflict situations is to display, assess, and retreat if necessary; i.e., no pure hawks or pure doves. 2. WAR OF ATTRITION MODEL - how long should you keep going Complex mathematical model, but the basic point is to figure out how long individuals should display aggressively before moving on! - e.g., Cats in a ‘stand off’ - How long to stay (and miss out on other opportunities) before giving up? 3. Sequential Assessment - making multiple different assessments along the way Basically, individuals assess their opponent’s fighting abilities over a number of bouts, increasing the level of aggression over the bouts. - e.g., Fish example: Start out by changing colour (visual assessment), then escalate to more dangerous behaviours (tail beating, wrestling, and circling/biting). OBVIOUS IMPLICATIONS FOR HUMAN CONFLICT Unlike nonhuman animals, we have access to weapons, so we can quickly escalate conflicts beyond threats, displays, or contests… We can also learn to become more aggressive. - Stable societies or communities have lower levels of aggression. Social disorganization/xenophobia can produce aggression. - Xenophobia - fear of others (us vs them mentality) …We could learn a lesson from bonobos! - when faced with conflict they will have sex CHAPTER 16 **PLAY One common characteristic of juvenile development in a wide range of species (e.g., mammals, birds, reptiles) is PLAY. Frequency of plaay is much higher in juveniles of every species than in adults. Defining plaay is difficult but we all know it when we see it! DEFINING PLAY - Plaay is a relatively new topic in animal behaviour research. It was previously considered the ‘garbage can area’: If you couldn’t categorize a behaviour as something else, people called it ‘play’! Play is very difficult to study experimentally (e.g., How do you ‘induce’ playing!?) ONE DEFINITION: “Play is a motor activity(somthng observed) performed postnatally that appears to be purposeless, in which motor patterns from other contexts may often be used in modified forms and altered temporal sequencing. If the activity is directed toward another living being it is called social play.” - Marc Bekoff and John Byers (1981) (IT LOOKS SIMILAR TO OTHER BHAVIOURS) Not a very satisfying definition, but it is very difficult to put into words, as you may have found doing your ethograms! **TYPES OF PLAY 1. OBJECT PLAY: Using inanimate objects (stick, rocks, branches, leaves, human toys, etc.; pushing, throwing, tearing, manipulating, etc.) - Quite common, and also seen in captivity where we provide animals with “enrichment” in the form of toys and other objects (e.g., lion - soccer ). Seen in lots of species, and often in birds like crows and ravens, and keas. 2. LOCOMOTOR-ROTATIONAL PLAY: Leaping, jumping, running, rolling, somersaulting, trampolining, etc. - Probably the most common type of plaay, seen in a wide variety of species! 3. SOCIAL PLAY: Plaaying with others Often seen in more ‘cognitively complex’ animals, like mammals - Example: Gorillas playing tag, kittens play fighting Also seen in other social animals, like parrots In zoos, we use the frequency and type of play as a measure of psychological well being; often provide objects to play with, even iPads. - e.g., We use computers and iPads at the zoo with the orangutans! Play can also be used to measure environmental stress in the wild! PLAY VS. AGGRESSION? Play often involves exaggerated movements, different vocalizations, or is performed more slowly than regular behaviours (Video: dogs https://www.youtube.com/watch?v=cSd5gYfRgoY&list=PLbPhJdEXzeOg53dZfGkFvbsVV2oxoYm0T) Some species have “PLAY MARKERS” to show conspecifics that they are just playing! PLAY BOWLING is META-COMMUNICATION (i.e., invitation to play - dog bow). Other “meta-communication” cues: - e.g., Open mouth play face e.g., Role reversal/self-handicapping ***FUNCTIONS OF PLAY ***Object play: Identifying new food sources (e.g., ravens) or learning how to be an effective predator (e.g., cat species). ***Locomotor play: Physiological benefits like increased motor skills and coordination, endurance and strength. ***Social play: Preparation for adult life (dominance relationships, testing themselves and conspecifics, developmental of social and cognitive skills). Generally, “Play allows animals to develop the physical and psychological skills they need to handle unexpected events in which they experience a loss of control (like being chased by predators, or being in a fight.” MECHANISMS OF PLAY Can be triggered by ENDOCRINE HORMONES, like testosterone. - e.g., More sex-based play in males than females, after puberty; Manipulate hormone levels to increase or decrease play! NEUROLOGICAL BASES: Dopamine tends to increase when animals anticipate play; serotonin and other neurotransmitters increase during play Play frequency and type can be manipulated genetically! - We artificially select for playfulness in some animals, like dogs, who retain their playfulness into adulthood (NEOTENY - infant like characteristics.. E.g. adult dogs play much more than adult wolves, along with bigger cuter eyes) WHAT SPECIES PLAY? Play is seen in a wide variety of species, but not all types of play are seen! Forms of play: SIMPLE PLAY ------------------→ COMPLEX PLAY Evolutionarily ancient animals ------------------→ Evolutionarily recent animals ^(Sharks, turtles, frogs, lizard)^ Evo ancient - simplistic pl@y Evo recent animals - complex pl@y Chapter 17 PERSONALITY ^(birds, marsupials, mammals)^ Studying personality in nonhuman animals is still controversial and in the beginning stages of research. It is important for you to know that Animal Behaviour as a field is still evolving - new topics are being explored all the time! Personality is a slightly ‘out there’ topic, so it’s fun to discuss! DEFINING ********PERSONALITY with animals (bold vs shy) - In humans, usually defined as “a pattern of RELATIVELY PERMANENT traits and unique characteristics that give both consistency and individuality to a person’s behavior” -Feist and Feist (2009) Personality traits that are inborn are often referred to as a person’s “temperament”. - The idea is that some traits are genetically determined. Other traits can be learned or shaped through experience. All of them, though, have to be a RELATIVELY considered part of a person’s persnality. PERMANENT PATTERN to be Studying personality in humans means to study individual differences in different traits. How does that translate when we are talking about animals? WE DON’T USE THE WORD PERSONALITY WITH ANIMALS AND USE “TEMPERMENT” INSTEAD - or “coping styles” or “alternative strategies” or “individual differences” rather than “persnality”. If you have had more than one pet, you’ve probably seen differences between them; you probably noticed differences in animals at the zoo as well. e.g., polar bears, orangutans, monkeys, lions, etc. THE PROBLEM: How do we distinguish personality differences from behavioural differences that are caused by the situation the animal is in? - We need to see long-term patterns in behaviour, which can take many years to observe! TYPES OF PERSONALITY TRAITS 1. In humans, a common personality trait continuum is extraversion/introversion. - Extroverts are typically: sociable, assertive, talkative and excitable Introverts are typically: quiet, reserved, less involved in social activities We all have a combination of these traits, (we are somewhere in between) but people tend to cluster toward one end of the continuum. This type of trait translates reasonably well into nonhuman behaviour… - Producers vs. scroungers (Foraging chapter) Some animals are *********BOLD’ and some are *******SHY in situations where novel objects are present; bold animals will approach, and shy individuals will retreat Bold fish tend to investigate new food sources and potential predators. Obviously, this can lead to some problems, because these kinds of behaviours can be risky and lead to death… Shy fish tend to NOT investigate potential predators or food sources, which is less risky but may mean they miss out on opportunities. You can see how these ‘persnality’ traits (or “ALTERNATIVE STRATEGIES”) would evolve in a species. - Returning to Game Theory (Remember hawks and doves?), boldness and shyness would be present in members of the population and would reach an equilibrium over evolutionary time. Overtime it equals out.. (the middle ground) Other traits studied in animals… - Sociability, assertiveness, curiosity, “agreeableness” (usually directed toward humans). Study on hyenas found reliable differences in all of these traits across individuals, not related to age/sex, although females were more assertive (as you would expect in hyenas!). These kinds of traits are also seen in almost every primate species. These innate differences all tend to come down to two main strategies through: Bold/shy = riskseeking/risk-averse, curious/not, fast/slow, independent/satellite, active/inactive all describe similar behavioural approaches or COPING STYLES. - **Satellite = nonterritorial ******* COPING STYLES: *****PROACTIVE: Characterized by aggression and territorial control in stressful situations *****REACTIVE: Characterized by withdrawal or immobility, and low levels of aggression - Proactive rats will attempt to bury or remove anything that causes them pain. Reactive rats tend to freeze or avoid novel stimuli that cause them pain These types of coping styles are seen in a number of different mammal species, and there are probably genetic and endocrinological (hormonal) mechanisms underlying the two COPING styles. EXAMPLES: Ruff birds: - Males “advertise” to attract females. Male ruffs are either bold (independent territory holders) or shy (non-territorial ‘satellite’ males) – genetically-determined. Females also carry the gene but it is not expressed in them! But if you give the females testosterone, suddenly they become independent vs. satellite as well. Great tit birds: - Males AND females are either fast (e.g. ,“bold”) or slow (e.g., “shy”). ”Slow” birds also take longer to explore their environment, and don’t benefit from social learning whereas “fast” birds do. Reproductive success is better when male/female personality types are the same! DO INVERTEBRATES HAVE “PERSONALITIES”? We know that octopus and squid are highly intelligent animals…but do they have ‘personalities”? Show differences in individuals in bold/shyness, activity levels, ‘anxiety’, as measured by distinct behavioural differences. We need to show relatively permanent patterns of behaviour and more recent research suggests this is not the case in octopus! But other invertebrate animals, like water striders, may have relatively permanent patterns of behaviour! These conclusions illustrate the importance of careful, long-term observations before any conclusions can be drawn. SO…WHAT’S THE POINT!? Is there anything to be gained from studying personality in animals?? Individual differences are REAL and can affect how well animals deal with STRESS! - e.g., Differences in orphan chimps and “rough scratching” (starting down by knees and then up into the armpits.. indicates they are stressed out.. it shows big individuals differences e.g., Differences in captivity/constant environment (e.g., orangutans, elephants, etc.) e.g., Differences in dog behaviour e.g., Differences in studying domestic animals, like pigs. https://www.youtube.com/watch?v=jmDlWdfe-sk ^^^ Variance in individuals can be applied to how they deal with stress differently
