Animal Behavior Exam 1
Why are bats using high frequency sounds in echolocation cries?
Bats make use of high frequency sounds For an object to reflect sound waves it must be larger than about 1/10 of the wavelength of the signal Therefore, high frequency sounds reflect from small objects better than lower frequency sounds do bats are pursuing small objects (like insects), so they have to rely on high frequency sounds
Ways of employing evolutionary theory in the study of behavior (in Comparative Psychology) problems with this
Comparative Psychology was still interested in what Romanes tried to do: building phylogenies for complex behaviors BUT.. the Historical (phylogenetic) approach—oldest and least powerful use of evolution to study behavior Problematic because: 1) Behavior doesn't fossilize (usually. footprints do) 2) Extant species used to represent ancestral types (representative species) -example: Rats representing mammals and pigeons representing birds. this is problematic bc for example, birds differ remarkably. so pigeons can definitely not represent them accurately 3) Separates a trait from others that, together, are the result of evolution to particular habitats -Hard to separate one trait from all others. -example: eyes don't evolve independent of the rest of the body. you can't look at them independently. same is true of learning. you can't understand learning by taking it apart from the constellation of other traits that have co-evolved with it. 4) Issues of convergent evolution -Convergent evolution: organisms that aren't closely related to evolve similar traits as they adapt to similar environments -Superficial appearance can't be the basis for putting a particular trait on a phylogenetic scale -example: camouflage on caterpillars is poisonous. on leaf insects it isn't. so how could you categorize these things together? you couldn't. they're not the same and they did not evolve for the same reason or in the same way. -another example: to really understand owl eyes, you would have to understand its lifestyle 4) didntrecognize importantance of genetics--- didnt believe it effected learning 5)believed in linear animal inteligence 6) believed lab work was superior to field work
Edward Thorndike
Comparative psychology developed way to assess animal learning Puzzle-Box -Put a cat in cage, and it can get itself out if it figures it out -Took less time over course of trials Also tested animals in mazes
Donald Griffin
Curvier concluded that bats use the "organs of touch" for orientation. but he was wrong •Griffin was an undergraduate at Harvard University • Coined the term "echolocation" • Later became the father of cognitive ethology -re-awakened interest in the animal mind.
Timeline of Field
Darwin (1859, 1872 )--> Romanes(1880s)--> Morgan (1890s-1900)--> Comparative Psychology and Ethology (1900-1960s)--> modern animal behavior studies
Order
Darwin (1860-90s) Romanes (1880s) Morgan (late 1890s) SPLIT (1960) Comparative psychology vs ethology Modern animal behvioral studies (1975 make up)
Objections to Darwin's Theory
Darwin did not anticipate the success of his book with general readers. Also did not anticipate negative reaction from science world 1) F. Jenkin criticized him for problems with BLENDING INHERITANCE/PAINTPOT PROBLEM. But, particulate inheritance! 2) Lord Kelvin—age of the earth calculations weren't consistent with Darwin 3) How could the earliest incipient stages of complex structures (e.g. eyes) have had adaptive function? 4) Incomplete fossil record
George John Romanes
Darwin's protégé Animal Intelligence (1882) Mental Evolution in Animals (1884) wanted to create a PHYLOGENY OF INTELLIGENCE. document evolution of intelligence. Organize from wholly unintelligent-->the most intelligent -first use of evolutionary theory to try to understand animal behavior. phylogenetic approach. behaviors, not only intelligence ANTHROPOMORPHIC/CENTRIC
Charles Lyell's
Darwins boss uniformitarian geology,finding new evidence that geological changes were slowand gradual rather than catastrophic.
Males and females anolis reproduction
Effect of castration and testosterone replacement on male Anolis courtship behavior -Testosterone will dictate actions. if you castrate a male, sexual interest goes away Anolis males -Dewlap (throat sack) -2 penises! Hemipenis Anolis Females -Reproductive cycle beings in spring -10-14-day cycle of receptivity linked to development of mature egg in an ovary -Ovaries produce increasing amounts of estrogen as egg develop -Estrogen travels via bloodstream to brain where it affects behavior: she becomes responsive to male courtship -Within 5-7 minutes after copulation female receptivity ceases due to production of prostaglandin—another hormone, stimulated by copulation -Mating only at the times when it is adaptive to do so!!!!!! REGULATION OF BEHAVIOR: hormones ensure that anolis are only reproducing when it is appropriate
Isolation Experiments
Eibl-Eibesfeldt squirrel study Ethology's attempt to show that FAPs were innate rear a squirrel in an isolated environment. deprive it of all experiences that might make them learn, and see what they still have Never gets opportunity to see other squirrel do this, no opportunity to dig, to de-shell nuts, etc. Later gave squirrel opportunity and it was able to! BUT..... Comparative psychologists argue with these types of studies. Daniel Lehrman is there anything you're missing that's remained in that environment? nature and nurture are not really separable. You must consider both
toad's visual system (who studied?)
Ewert small, moving object (w/ shape/worm vs antiworm movement-axis) in the excitatory region -------------> stimulate gangleion cell thalamus (brain region) - "predator detector" -not only fires, also limits optic tectum firing (predator>prey) optic tectum (brain region) - "worm detector" illustrates umvelt--> N.S. shaped to discriminate stimuli -Peripheral ♣ Retina with rods and cones ♣ Areas of responsibility—individual bipolar cells ♣ Inhibitory and excitatory cells ♣ Ganglia cells control input to bipolar cells ♣ Receptive field of ganglion cell controls optic nerves unless the worm moves, ganglion cell will NOT fire. this is called STIMULUS FILTERING
How could the earliest incipient stages of complex structures (e.g. eyes) have had adaptive function?
Ex. The eye—doesn't it have to be working in its complexity in order for it to do its job? Darwin said you can start simple, in a way that doesn't reflect in the manner of which the trait will ultimately be adaptive Example: Extant mollusks • Different visual systems associated with lifestyles • Clam doesn't have sophisticated eye, but it doesn't need one for its lifestyle • Original advantage can be very different, and given sufficient evolutionary time, that additional complexity will evolve -contexts change, environments change
Exposure to stimuli before hatching
Exposure to stimuli before hatching influences imprinting!!!!! woman talks to egg--something for chick to orient to when it hatches then it hatches and imprints on her hatching synchrony!!!! (see next card)
Blending inheritance and the Paintpot Problem
F. Jenkin criticism of Darwin The Paintpot Problem: -Blending inheritance: offspring are a blend of the parents. The blend of characteristics is passed on to the next generation -Jenkin said variation would be washed out over time if this were true. (Red/white card example. With mating over time, you would just get a diluted version of the original characteristic. Individual redness would get lost over timebenefits would be eventually gone) -BUT...Gregor Mendel and Particulate inheritance: offspring are a combination of the parents. Parental contributions are passed to the next generation as separate entities. Variation is maintained over time. -Mendel said that traits are inherited as "particles". The answer to the problem! Genes. -things don't blend. they're passed down separately. a tall parent + short parent does not equal a medium child. will be either tall or short! no blending. alleles.
FAP
Fixed Action Pattern (F.A.P.) - The unit of inherited behavior - goes to COMPLETION even if stimulis (or egg) is removed Not a reflex!!!!! Behavior occurring at appropriate times. Regulation of behavior. Serves a purpose/is adaptive presence of hormones
Incomplete fossil record
Fossil finds: • Archaeopteryx • Xiaotingia—first bird • Changes in cranial shape in the horse lineage (Hyracotherium--better known as eohippus ("the dawn horse").) -Darwin thought fossil findings will occur when conditions are right. He never thought there would be a huge volume of evidence. so this wasn't a problem for him. in time , more and more evidence would be discovered -The human family tree: recent fossils found in Ethiopia in the last 10 years include Ardipithecus kadabba bones from over five million years ago, left, teeth of the Australopithecus anamensis, which lived four million years ago; and the skull and lower limbs of a 3-year-old Australopithecus afarensis, thought to be 3.3 million years old. At right, teeth of the afarensis, anamensis and the modern chimpanzee. -more and more evidence discovered over time
Historical (phylogenic) approach
George John Romanes -first use of evolutionary theory to try to understand animal behavior. phylogenetic approach. behaviors, not only intelligence -attempts to find "behavioral homologies" oldest and least powerful use of evolution to study behavior
hatching synchrony studies
Gotleib 1965 1) remove eggs from mother, put them in separate incubators control their auditory experience -Can control and record vocalizations produced by gosling (either their mom or their siblings) -If you deny them these vocalizations, they won't synchronize 2) -deprive mallard duck of any vocalizations -then test their willingness to follow a model (stuffed duck) -play vocalizations of mallard duck, wood duck, chicken, and silence -there is a preference to follow your own species' sound! -this suggests that not all auditory stimuli are equal. there is a bias towards your own species' sound But is this an INNATE preference? ....... Later studies show NO! - they loose this bias if you mute them surgically. - so it has to do with the vocalizations the duck makes within the egg! - so if they have neither heard a sound in egg, nor made their own they follow whatever they find post hatching
FAP Examples
Greylag goose female adjusts herself on top of the egg/moves it around and always retrieves an egg that has been bumped out of her nest in the same manner This is a fixed action pattern She carries this sequence to the completion, even if the egg slips away during the process As if it were a unit of behavior from start to finish Rolls egg back, even if you remove the egg, she will continue this action as if the egg was there. Will go to completion even if immediate stimuli is not there Not a reflex. Behavior occurring at appropriate times. Regulation of behavior. Serves a purpose/is adaptive Restrictive to breeding season: Response occurs during nesting and incubation of eggs. (Presence of hormones) There will be a refractory period after an FAP occurrence during which animal is less responsive to releaser
Salmon Experiment, Hasler
Hasler reared hatchlings in tanks with non-organic "odors" -hat chlings marked and released - "a form of olfactory imprinting" - these salmon returned to streamsbaited with this chemical
hormones
Hormones are substances secreted in one part of the body that cause changes in other parts of the body. Source: endocrine glands or neurons ORGANIZATIONAL effects - Occur during development. Early in life - just before or right after birth - Tend to be permanent----> effect NS layout/development ACTIVATIONAL effects - Usually occur in adulthood - Tend to be transient (last only as long as the hormone is present)
mouse hormones in uterus study (whose study?)
Hormones: organizational effects vom Saal & Bronson Shows horomone response VARIATION WITHIN SPECIES (vs birds btwn species) within uterus, males and females next to each other (siblings)—will they be influenced by being surrounded by those of different sex -Looking at what hormonal differences will have on behavior -a female developing next to 2 males might be impacted by the T. Developmental exposure will influence behavior in a dramatic way -Example: 2 m females (have brothers developing next to them) and 0 m females (no brothers) -The difference in hormonal exposure is only happening before birth, so whatever behavioral consequences occur is a result of this early exposure (so these are ORGANIZATIONAL effects!!!!) - miniscule amounts for short time and yet longlasting effects -Organizational effect—hormones wont be there anymore, but persistent RESULTS: see other slides
Vestiges of the Natural History of Creation Robert Chambers
In 1844, a popular book, led middle-class readersto accept that the progressive evolution of life was part ofthe divine plan of creation.
I.R.M
Innate Releasing Mechanism (I.R.M.) -"black box approach": Something that we know is important and accept that its relevant, but we don't really have understanding of what's going on at this time - Decide, system or object which can be viewed in terms of its input, output, and transfer of characteristics without any knowledge of its internal workings -Simple nervous system "gate keeper" • DETECTS key stimulus, and GOVERNS performance of behavior
Migration
Involves navigation: periodic movement from one location to another. seasonal
Homing
Involves navigation: the ability to return to a home site or locale after being displaced. independent of experience!!! doesn't even have to be familiar with the environment!
tropisms
J.Loeb plant and animal behavior can be explained by physiochemical reactions to stimuli (tropisms). not consciousness
Simple orientation mechanisms
KINESIS: Non-directed movement in response to environmental factors such as light, temperature, moisture (e.g., pill-bugs) Terrestrial Isopods in the Class Crustacea. Need relative humidity about 70%. (Crustaceans include lobsters, crabs, shrimp). pill bugs are related to crustaceans—When they get exposed to a change in humidity, they begin moving randomly. As the relative humidity increases and conditions become more desirable, their movements begin to slow down. This movement is RANDOM!!!!! Not directed, just a response. They are not tracking the humidity TAXIS: Directed movement; the source or a gradient of some factor are detected and the organism moves in the appropriate direction (e.g., tick) ex: "questing" tick will position itself at top of blade of grass bc it responds to pull of gravity ex: male orients towards female from smell, then can find the female
Imprinting and Lorenz's conclusions about it
LORENZ important to ethology A form of early learning in which the young animal forms an association or attachment with another animal (object, or class of items in the lab.) • Early conclusions of LORENZ 1) Many species, especially precocial ones, showed a marked critical period during which exposure to the imprinting objects must occur. 2) The strength of the imprinting depends on the effort expended by the animal in following the object. 3) Imprinting was irreversible. 4) Animals could be imprinted equally well to virtually any object or organism. TESTED THIS ON A RED SQUARE IN THE LAB. 5) Animals learned the species identity of their sexual partners at the same time that they imprinted on their mothers.
Vacuum activities and the hydraulic model
LORENZ sometimes, FAPs occurred when no releaser was present (e.g., in zoos): vacuum activities Why? Lorenz proposed his hydraulic model of action specific energy Every FAP would have its own action specific energy. Within organism, there is a reservoir for this specific energy. energy in the reservoir would build up. in ordinary circumstances in nature, a releaser would be encountered and then the action specific energy would be used to fuel the occurrence of the FAP. when there is no releaser (in zoos), the reservoir just gets to the point where it can no longer be contained. action specific energy will spill out-->FAP will occur even though there is no releaser critics said there is no evidence for this.
Cues for navigation
Landmarks Magnetic Fields - Pigeon (backup mechanism) - Sea turtles Chemical Cues / Olfaction - Salmon Sound Cues - Ducklings orient to maternal calls - Bats: echolocation Electric Cues - Electric fishes Ocean Currents - Lobsters - Sea Turtles - - - Sun - Tinbergen's digger wasps Bees Arctic tern migration (coastline) Bees, ants - Stars Birds (e.g., pigeons) - Birds (indigo bunting) (planetarium studies) Lunar / Tidal - Fiddler crabs - Grunions
Baby Schema (who?)
Lorenz said this is an example of FAPs in humans! what we find cute in babies: big head and big eyes these are like releasers for us Head and eyes large in proportion to body, rounded body shape Teddy bears capture and enhance those features and make them "cuter" Speaks to evolution of cartoon characters • Evolved to be cuter as a result of how the audience reacted
NAMES and WHO ARE THEY************
Lorenz: Ethology -- animal mating Tinbergen: Ethology Spallanzani: Blind vs deafbats
Comparative Psychology
Morgan-->Comp Psych 1900 to early 1960's More careful studies could be done in lab where variables can be controlled and experiments conducted with rigor. -Desire to go along with Morgan -Careful studies where you can control variables Learning easily studied in the lab: learning apparatus. -A place where you can exercise a high level of control and rigor -If you start with this more controlled situation, you are looking at a small variety of animals. you can't study elephants in a lab! they were still kind of interested in what Romanes tried to do: A major focus was the construction of behavioral phylogenies, especially LEARNING (Wanted to see how more complex forms of behavior might have evolved)
Experimental setup to study moth's auditory system
Moth auditory system can detect: 1) how close bat is (proximity) 2) what direction bat is coming from (direction) ________________________________________________________ ROEDER!!! Bombarded moth with sounds, to test its reaction to different kinds of auditory stimulation then they recorded the neural activity with electrodes found that the neurons were responding to sounds of bats' orienting cries two receptors in moth auditory system: A1 and A2 A1 receptor: -low intensity stimulus-->neurons show some baseline activity - informs how close bat is (firing rate) -high intensity stimulus (similar to bat moving closer to the moth)-->A1 fires more. -fires more as sound gets louder! so moth has ability to detect proximity of bat -if bat approaches from the left, there will be a slight difference in firing. so it can also detect direction A2 receptor: -A2 doesn't fire more than baseline, even when bat gets closer -it only fires when the highest level of amplitude happens (when the bat is as close as it could be). when this happens, the moth responds by shutting down its flight system! so then the moth is much more difficult to pursue because of its unpredictable flight pattern
Noctuid moth auditory system
Moths are frequent prey of bats Some moths have evolved ears of their own that help them deal with bat predation(increase survival 40%) - detects cry of batbeforebat registers echo -PRIMARY defenses prevent detection by predator, SECONDARY defenses enhance survival after detection - moth turning away from bat prior to detection = priamry defense - complex aerobatic maneuvers or sound productionafter detection = secondary defenses they can detect the presence of bat and make it less likely that they will be found VERY SIMPLE auditory system, but very effective
Map sense vs. compass sense
NAVIGATION map sense: if you were stranded at airport, you could get back to emory bc you have a mental map. map sense is made up of memorized landscape snapshots compass sense: you know you have togo northeast. but how do you know which direction is northeast? that's compass sense
Orientation
Organism positions itself in the environment on the basis of external cues.
What is it like to be a bat?
Paper by Thomas Nagel -consciousness and subjective experience -philosopher Nagel chose bats as an example of an animal with very different forms of perception, sensation, and "umwelt" compared to our own species -it's hard for us to even know how other humans see the world (umwelt)
Alternatives to Darwinian Evolution (Parson Naturalists)
Parson naturalists (documented stuff in nature with the goal of showing that the world was so complex and only God could have created it) -Scientific creationism, intelligent design (both trying to preserve idea of God. God controlled evolutionary process) 1) John Ray: The Wisdom of God Manifested in the Works of the Creation 2) William Paley: Natural Theology, Evidences of the Existence and Attributes of the Deity, Collected from the Appearances of Nature -Paley's argument: the watchmaker: The complexity of a watchthere must be a watchmaker (God). Must be an intelligent designer -Dawkins says the watch analogy is false. "Natural selection has no purpose in mind. It's like the blind watchmaker. This explanation is sufficient". -Ray and Paley argued that God created the world
Early Ethologists
Period from about the turn of the century to the early 1960's Early Ethologists: 1) Von Euxkull -Concept of the Umwelt: the world experienced by a particular organism -Animals have different sensory organs that evolved. so experience varies depending on the species and how they've evolved 2) Lorenz -geese, Ideas of imprinting, FAP and action sepcific actions 3) Tinbergen -Why particular behaviors would've been selected for 4) Von Frisch -communication in bees
"The Snark was a Boojum"
Poem by Lewis Carroll Frank Beach some kinds of snarks are dangerous in the poem. similarly, there are some things in comparative psychology that have developed that we need to be careful of You can't just study rats, and you can't just use lab and apparatuses, you have to expand this
Lord Kelvin's Criticism of Darwin
Questioned Darwin's thinking based on his calculations of the age of the earth Darwin argued that the world was far older than the bible did Lord Kelvin was very religious Presented an argue based on physics and math • Said that the earth's temperature increased one degree Fahrenheit for each 50 ft you went under ground. So he determined that the earth as 100 million years old Darwin argued that the earth was older and there was time for natural selections Kelvin didn't know about radioactive decay Later, we learned he was wrong about the temperature thing. Today, we know that Earth's temperature variation is affected by radioactive decay, and that Kelvin's cooling calculation was worthless for telling us the age of our planet.
Cockroach escape behavior
Roaches are very successful at evading threat (like your shoe). Cerci/cercus (appendages on the hind end of the roach) responds to WIND created by approach of a predator-->cockroach will turn and run away from threat. Simple LOCALIZED response, no categorization of threat orbrain usage
cataglyphis ant
Rüdiger Wehner and his studies of the desert ant, Cataglyphis: they know it uses sun/polarized light for navigaiton - dont know HOW it does it -but use odomoter form to measure distance (cutting leggs traveled 50% less vs lengthening travelled 50% more)
Bowler
Scientists didn't believe Darwin at first Darwin was criticized for hiding the fact that he was not the first person to propose a theory of evolution ancient Greeks had theories But the philosophers Plato and Aristotle both argued that the form of each species of life was absolutely fixed BELIEFS ABOUT GOD AND ADAPTATION: The study of living things confirmed that each species was carefully adapted to a particular way of life, thereby displaying the Creator's wisdom and benevolence. This was the 'argument from design', intended to prove the existence of a wise and benevolent Creator from the observed nature of His handiwork. In 1844, the Edinburgh publisher Robert Chambers wrote a popular book, Vestiges of the Natural History of Creation, which led middle-class readers to accept that the progressive evolution of life was part of the divine plan of creation On the HMS Beagle, Darwin was converted to Charles Lyell's uniformitarian geology, finding new evidence that geological changes were slow and gradual rather than catastrophic. Darwin also collected fossils and studied the geographical distribution of species. It was on the Gala ́ pagos Islands that he found the evidence which converted him to evolution. The various islands were inhabited by distinct species of finches, which had almost certainly evolved from small populations originally derived from a single original species. Darwin was soon convinced that isolated popula- tions evolved to adapt to their local environment. General- izing from this, he conceived evolution as an ever- branching tree, some branches becoming extinct while others subdivide and move off on different tracks. Darwin was impressed by Thomas Malthus' 'principle of population', according to which the population always tends to outstrip the food supply. This leads to a 'struggle for existence' Darwin developed his theory in the late 1830s but did not publish. In part, this was because he was afraid of public controversy. But he also realized that he still had much to do if his theory was to answer the questions that would be raised by other naturalists. In the mid-1850s Darwin began to write up his theory for publication. He was interrupted in 1858 by the arrival of a paper by Alfred Russel Wallace outlining what appeared to be a theory very close to his own. There has been much controversy over the relationship between Darwin and Wallace, some partisans of the latter accusing Darwin of deliberately sidelining the independent discovery. Others note significant differences between Wallace's outline and Darwin's theory We are told that the one factor that held back acceptance of natural selection was his failure to appreciate Mendel's idea that heredity is particulate (Darwin) fossils provided evidence of evolution for people heredity had also become a focus of attention for Darwin and the relatively small number of biologists who took natural selection seriously from the start. Darwin himself was concerned when Fleeming Jenkin argued in 1867 that natural selection could not work if heredity was a process that blended the characters of the two parents together. The effect of a single favourable variation (a sport) would soon be diluted by breeding with unchanged individuals.
bat sound waves
Sound waves exist as variations of pressure in a medium such as air. They are created by the vibration of an object which then causes the air surrounding it to vibrate. Wave length and amplitude (loudness) hertz (Hz): the number of cycles occurring per second --bats make use of high frequency sounds There are more cycles in the same period of time in a high-pitch sound than in a low-pitch sound. Thus, high-pitch sounds such as a bat's echolocation cry provide more information to accurately represent than do lower pitched sounds
Spallanzani
Spallanzani • Noted owls could not fly in complete darkness, but bats could • Blinded bats returned to their roost with full stomachs • Deafened bats (plugged their ears with wax) did not navigate or feed proficiently
mouse hormones in uterus study RESULTS
Subsequent adult behavior: females Results of prenatal exposure to testosterone - compared to 0m adult females, adult 2M females: - Are less attractive to males - Are more aggressive to female intruders - Scent-mark (with urine) a novel environment at higher rates - Have longer and more irregular estrus cycles Note: After birth the testosterone levels in all female mice were similar(shows that it is organizational, not activational) Subsequent adult behavior: males • How does differential exposure to estrogen affect male mice? - 2m males exhibit parental behavior when presented with a newborn pup. - 0m males usually kill newborn pups. -counterintuitive!!! shows that you can't predict what kind of behavior a hormone will produce.
Super Releasers
Super releasers take priority these super releasers seem to take priority over the real thing. How does this make sense evolutionarily? There was never the need to discriminate between huge egg and small egg, so bird doesn't know how. example: red pencil example: when multiple eggs roll out from under birds, the bird will go for the bigger one (even if it is much bigger than their typical egg) example: Australian male JEWEL beetles will try to copulate with discarded beer bottles, but they have to be of the right type, brown ones with a dimpled surface -Super releaser -Will die in sun before leaving example: ash tree population has been devastated by EABs (emerald ash borers) beetles. Ash tree bark appears purple. So they used a "barney trap"—purple thing in tree. Super releaser, so the beetles just stay on that instead of the actual bark-->don't ruin the bark
Gulls releasers
TINBERGEN Red spot on gull bills Gull chicks peck on the red spot (feeding response). Tinbergen used models to study the releasers involved: some with red dot, some without, some without correct shape, etc. (slide 7. Take note of bar D, in which chicks respond to red pencil. Interesting. Shows that releaser=red) • How do we determine what the key releaser is? o Simple models: gulls head, control shape, color of dot and compared responsiveness of chicks when presented with different models o A) Representation of head and red dot ♣ Second most effective o B) Preserves bill and red dot but no head ♣ Third most effective o C) Head but no red dot ♣ Least effective o D) Red pencil ♣ Most effective -does the position of the red dot matter? ...configuration of stimuli -->questions about the IRM! concluded that releasers and the IRM were more complicated...perhaps the nervous system comes with inherited "images" of the correct configuration? -dot must be in the right place on the bill in order to get the largest response -if the image doesn't match the internal image, then the response will be less -BUT...replication by Jack HAILMAN showed this was wrong
Importance of testing evaluative hypotheses
TINBERGEN emphasized this idea. Have to test with data, not just come up with a conclusion! Problem of pure speculation about adaptive significance 1) Tinbergen tested 3 different hypotheses for the egg shell problem. 2) Why are flamingos pink? -camouflage to avoid predation when the sun sets or rises? Pink color allows them to blend in at sunrise/sunset. -nope! It does benefit them, but it is actually just due to pigmentation of their food! -Epiphenomenon: it's a secondary issue. it's not necessarily adaptive, just occurs from their habits
Male stickleback fish releasers
TINBERGEN!! -Male will establish territory, build a nest, and protect it -Another male coming will lead it to attack -Tinbergen had a Stickleback aquarium -Noticed male demonstrating aggressive behaviors, looked outside and saw red mail truck -Why would he confuse this for another fish? Red color cue --> Tinbergen used simple models -First one: looks like stickleback but not red. Elicits no response -when there is red, almost instantaneous response -Third: doesn't look much like a fish, but is red. Almost instantaneous response
toad's visual system cont'd
Thalamus and optic tectum Worm detector cells in the optic tectum: fire when there is the right shape and right movement Thalamus -Cells in here responding to the presence of a predator -when they detect a predator, they inhibit optic tectum, shutting it down. so it won't worry about worms now. escaping predator is first priority
egg study, caswell
The length of an egg correlates with bird body size. The shape of an egg—how asymmetrical or elliptical it is—relates to flying habits. And the stronger a bird's flight, the more asymmetrical or elliptical its eggs will be.
Regulation of behavior by hormones: adaptive responses
The neural and hormonal mechanisms that underpin reproduction can be viewed as adaptive responses that restrict this behavior to times when favorable environmental conditions are likely. ANOLIS example. they only reproduce in the spring
Refractory period
There will be a refractory period after an FAP occurrence during which animal is less responsive to releaser if releaser is presented repeatedly
Tinbergen's "4 whys"
Tinbergen upset that there was so much disagreement between comp psych and ethology. he came up with these 4 questions to show that you could approach things in different ways. To fully understand behavior, we need to ask questions about: 1) Survival value or function 2) Causation 3) Development/Ontogeny 4) Evolutionary history/Phylogeny ex. 1. injury/diseas to chicks? effects predation? 2. why does this cause response? (releaser white color --- hormones w incubation/nesting) 3. FAP but some learning--- 1st breaders will remove shells even before laying eggs...... but birds preferentially romve dummy shells of same color as fake eggs they incubate... learned "fine tuning" response to innate egg removal 4. construct historic phylogeny of egg shell removal/not removal based on extant species (ex kittiwake gulls)
study of landmark usage by digger wasps (Who?)
Tinbergen's !!!!!! an example of learning -Female digger wasps dig a hole, go to get prey, and comes back with prey and puts it in hole -before she leaves, she flies around nest -Why does she fly around nest? -Maybe she was surveying local conditions (landmarks and stuff) so that she can return more easily -Tinbergen put pinecones around the hole, then moves them before she returns. -she still flies inside of the pinecone circle, even though it's in a different place now. she is making use of the landmarks
Proximate vs. ultimate factors
ULTIMATE: ultimate causation explains traits in terms of evolutionary forces acting on them -Survival value -Why did the ancestors show behaviors that are present today? -Helped them stay alive, reproduce PROXIMATE: proximate causation explains biological function in terms of immediate physiological or environmental factors
"Principle of serviceable associated habits" and "Principle of antithesis"
Variation of Animals and Plants under Domestication (1868). The Descent of Man (1871) Expression of the Emotions in Man and Animals (1872)**** -interested in internal, emotional states - "Principle of serviceable associated habits": particular state of mind-->same behavior. example: a dog will circle around before it lies down to take a nap. ancestrally, this was helpful to tramp down vegetation, mark territory, scatter ticks, etc. not helpful anymore, but behavior still preserved! - "Principle of antithesis": when animals benefit from communicating about their internal states, they express different states very differently. to make sure that ppl won't get confused. Darwin's dog Polly example. the way they behave is shaped by natural selection o Ex. Aggression vs. submission ♣ Aggression: tail up, hair up ♣ Submission: Arched back, tail down
Evaluative approach
Way of Employing Evolutionary Theory Ethology This approach seeks to evaluate a behavior by how it contributes to the fitness of animals performing it. The rationale is that behaviors that have been selected for must be advantageous. What is the nature of that advantage? ADAPTIVE SIGNIFICANCE of things. 1) EX: Tinbergen's evaluation of eggshell removal by gulls after chick hatches, bird takes egg shell and flies it away. what benefit does this have? TINBERGEN tested 3 different hypotheses: 1) egg shells could cause injury to chicks? (bc jagged edges and chicks are clumsy). 2) getting rid of egg shell prevents disease? Bc bacteria can grow in egg shell 3) getting rid of egg shell reduces predation? Eggs are easily spotted (color and jaggedness of broken shell), might attract predators -he painted chicken eggs so that they would resemble gull eggs. then put them near nests. RESULTS: predation increased when broken egg shell was nearby 2) Adaptive value of mobbing in gulls -Kruuk -Mobbing behavior in Black-headed gull colony. they collectively mob an intruder -How do we test that mobbing behavior deters predators? -Moved eggs around and inside of colony, tested them at different points. -Results: risk is least great to eggs within the depths of the colony, intermediate on border, outside of colony all taken by predators -so mobbing behavior is advantageous because it protects eggs from predators mobing behavior shows both divergent (kittiwake) and convergent evolution (Gulls and swallows both swarming)
Echolocation
You produce a sound and then it bounces backs and provides information about objects Echolocation involves the production of sound and the reception of echoes that return from the objects. By comparing the outgoing pulse with the returning echoes—which are modified and less powerful versions to the outgoing pulse—the bat's brain can process information
taxon
a particular class (class, genus, species)
locusts
african tribes take advantage of high frequency hearin gof locusts by rapping metallic pots to deflect swarms of locusts from their property
Jack Hailman
after the gull bill questions about IRM and configuration "Ontogeny of an instinct" replicated gull red spot experiment found that configuration was NOT important: the dot was equally effective no matter where it was on the model IF it was presented at the same rate of movement. also found that young chicks' pecking accuracy (hitting the target) improved over time. there is CHANGE in FAPs...learning! "ontogeny of an instinct"=development of an instinct. FAPs can develop/improve, even though they are innate -improvemtn could have been learning (via exposure) or possibly just eye development (sight improve) dropped the idea of an IRM, replaced it with the idea of feature detectors in the nervous system
ethogram
an formal description or inventory of an animal's behaviors
Negative results
an outcome in which the null hypothesis is not rejected, and thus the alternate hypothesis is rejected
4 whys example
asking the 4 questions about Eggshell removal 1) Survival value ♣ Reduces predation 2) Causation ♣ Releaser: white inner color; jagged edge ♣ Hormones associated with incubation and nesting 3) Development ♣ FAP, so even first time breeders will do this ♣ Birds given artificial eggs of unnatural colors to incubate, preferentially remove dummy shells of the same color. This seems to be a learned response "fine-tuning" the innate egg-removal behavior 4) Evolutionary history ♣ Phylogeny ♣ Possible that some species lost it ♣ There are species of gulls (ex kittiwake gulls) who don't remove eggshells because they tend to nest on cliffs, making them less vulnerable to many kinds of predators
clock shifting in pigeons
birds clock shifted in lab compensated for this "biological clock component" when they were released at 6am and tried to return back back up: orientation mechanism is magnetic field detection (ex on cloudy day) **clockshifted monarch butterflies behave similarly!!
Herbert Spencer
called the 'survival of thefittest'.
anatomy of a controversy
challenge to dance language - Wenner & Wells concerns - olfactory cues (pheromones) secreted by workers when they mark source of nectar/water - this unique scent is recognizable by bees of same colony - their studies shows 87%of recruits failed to arrive at feeding dish, those that did arrive took 30x longer than direct flight
taxonomy
classification of organisms
Sexual motivation in male guinea pigs: individual differences in response to T.
correlation and manipulation there are differences in male guinea pig sex drive castrate the males-->THOSE DIFERENCES COLLAPSE. after castration, all guinea pigs pretty much have the same level of sex drive then, replace hormone--add testosterone back (they all get the same level) even though they all RECIEVE the SAME level, the behavioral DIFFERENCES emerge again! some individuals are more RESPONSIVE to the T
Species differences in the effects of testosterone
correlational studies good example of the VARIATION that can occur both between and within species!!! there is no one pattern for the relationship between T and territoriality. sometimes territoriality surges along with a surge in T, but other times it is at its peak when there is no T at all looking at correlation between presence of testosterone and whether or not a territory is maintained and defended by birds -Lapland longspur: testosterone produced and territorial behavior is a tight fit -White crowned sparrow: territorial behavior that begins as testosterone peaks, but as testosterone decreases, it is maintained for a period of time and then goes away when T is gone -Puget sound white-crowned sparrow: Longer drop but similar -Song sparrow: behavior with testosterone and then goes away when it does, but comes back without testosterone -White-browed sparrow-weaver: only small amount of testosterone, territorial behavior still exists. so no correlation between T and territorial behavior • Some have two mating times • Even diff animals within a species can have difference associations between presence of hormones and particular outcomes.
extant
currently existing species
Ontogeny
development
J.Loeb
disagrees with the idea that plants have consciousness. says we don't need anthropomorphic interpretations he says "tropisms"-plant and animal behavior can be explained by physiochemical reactions to stimuli
map sense and compass sense in honeybees
distinguishing map sense and compass sense in honeybees: -compass sense comes from position of the sun -bees make use of landmarks to create map sense, but it's dependent on experience -Map sense is made up of memorized landscape snapshots STUDY: -place sugar water near hive, bee will travel to visit it -when it gets there, trap it and move it 400 meters east. then free it -will it be able to get back to the hive? -DEPENDS ON HOW EXPERIENCED THE BEES ARE! -experienced bees, who have foraged for a long time, know the landmarks so they can find their way back -naive bees dont take into account displacement -so...it depends on age and experience - u need experience for map sense
Key scientists in bat echolocation
donald griffin- basics of bat echolocation (blind vs deaf bats flying in dark) Roeder- a1 a2 neuron receptors initial hypothesis fullard- controversy with Roeder on specifics of a2
evolution controversy
evolutionary change does not mean increasing complexity or progress example: cavefish have no need to react to light-->they've evolved SIMPLE eyes. not structurally complex
non auditory defenses to bat predation
fly close to vegetation in "clutter zone" where echos of prey are lost in vegitation echos
optimality models
form of predictive approach - selection of genes related to efficency -tasks critical to survival/reproduction should be acted on by N.S. -"Animal economics" advantages: - testable, quantitaive predictions - assumptions are explicit (ex. starlings optimise rate of food delivery) disadvantages - if behavior doesnt fit predictions? - if behavior is not optimalized? -ex enviro change vs insufficent genetic variation - good enough vs optimal -assumptions of genes for behavior -constraints on optimality ..... ex JUNCO feeding!!
Gould
further tests of dance language 1st experiment - lock recuits in and replace initial sugar water to remove an yscent - place many sources around hive that are in diff directions + closer - release recuits + moniter behavior - majority flew farther to the waggle indicated source regardless that there were no olfactory cues - indicates that von frish hypothesis IS accurate about dance language 2nd experiment - misinformed bees w/ light into hive - bees give priority to light over the vertical revealing that misinforming is possible indicated that bees are paying attention to dance language - (light more deeply rooted) riley and collegues answered Wells& werner concerns shows: using very accurate microtrackers show recruits fly to VICINITY of food source, THEN search for location using odor, etc - more effective anyways salvage von frish hupothesis while enhancing understanding
oxytocin and male recognition of females in mice
gene manipulation present male with female if you present the male with the same female multiple times, the male eventually gets disinterested if you put a new female in, his interest rebounds knockout procedure: eliminate a gene that is critical to the production of oxytocin after the knockout-->male continues to show interest to the same female presented in successive trials conclusion: this is some sort of social amnesia. in some way, they aren't remembering their previous encounters with the female
O. C. Marsh
he fossil horses discovered by O. C. Marsh in America were hailed by Huxley as 'demonstrative evidence of evolution' because they showed the stages by which the specialized modern form had evolved from a generalized mammalian ancestor.
von frisch
honey bee dance language development - round dance if nearby source (no directional cues) waggle dance if source far from hive - number of waggles indicates distance, orientation of straight run waggle in relation to vertical indicates direction of source in relation to sun - this is adapting to hives in darkness (asian honey bee dances on flat surface + runs directly as food source) - it appears sun is more deeply rooted in bee understanding -when light is shown into hive blinded bee will do dance but recruits interpret dance in relation to light orientation rather than vertical
why do hormones effect different animals differently?
hormones impact how genes work
katydids
in response to echolocation of sympatric bats, they shut down their acoustic courtship displays to avoid passtive detection by gleaning bats
Additional adaptations for echolocation
inferior colliculus lateral lemniscus
Thomas Malthus'
influnced darwin Thomas Malthus' 'principle ofpopulation', according to which the population alwaystends to outstrip the food supply. This leads to a 'strugglefor existence' in which many must die
Ethology and instinct
instinct=inherited behavior Fixed Action Pattern (F.A.P.) - The unit of inherited behavior Releasers -(generally) requires releaser aka sign stimuli - the environmental stimulus the prompts the organism to perform the FAP -behavior occurs when it is adaptive/when the situation calls for it. not randomly - occurence leads to refractory period - threshold for response changes as fxn of internal conditions - releaser-response patterns conspecific - learning NOT NECESSARY but possible! -sensory systems have evolved to act in a certain way in terms of particular situations in the environment Innate Releasing Mechanism (I.R.M.) -"black box approach": Something that we know is important and accept that its relevant, but we don't really have understanding of what's going on at this time - Decide, system or object which can be viewed in terms of its input, output, and transfer of characteristics without any knowledge of its internal workings -??????? figure this out Ethology's attempt to show that FAPs were innate: Isolation Experiments -e.g., Eibl-Eibesfeldt squirrel study
animal behavior
internally coordinated but EXTERNALLY visible. not heart rate. has to be an actual behavior
Ethology's attempt to show that FAPs were innate
isolation experiments (squirrel)
Tracking neurons
lateral lemniscus- inhibit transmission of sound to higher regions of brain during echolocation + inner ear bone seperation inferior colliculus- "echo detector" cells sensitive to the SECOND of two sound stimuli aka sound bounce specialized Tracking neurons- present in both^^ they remain active so long as interval between cry + echo decreases In both areas these neurons remain active as long as the interval between the orienting cry and the echo continues to decrease (youre closer to the object--the interval decreasing means it's taking less time to come back to you) these neurons are providing information about the successful pursuit of your prey
What can we conclude from the isolation experiments?
nature and nurture are not really separable. You must consider both
Neuroethology
neuroscience of animal behavior Since the mid 1960's much research, inspired by the early ethologists, has focused attention on how the nervous system influences and helps regulate animal behavior -studied feature detectors in the nervous system (instead of IRM, started to understand more) -Feature detection: process by which the nervous system sorts or FILTERS complex natural stimuli extracting behaviorally RELEVENT cues that are associated with important objects or organisms in their environment, as opposed to irrelevant background or noise -Biological bases of the regulation of behavior -Specialization of the nervous system -Hormones
Modifications of early views of Lorenz on sexual imprinting
o Sexual imprinting and maternal imprinting don't occur at the same time in all species o Critical period not as restricted as originally thought o Not all objects/organisms are equally effective (example: exposure to the vocalizations causes a bias for certain objects in the environment) o Imprinting is reversible within the sensitive period, especially if initial object is not conspecific (member of your own species) -not true. the original bond can be broken, especially if it's not a member of your own species
Nature red in tooth and claw **on exam
poem by Alfred, Lord Tennyson A reference to the sometimes violent natural world, in which predatory animals unsentimentally cover their teeth and claws with the blood of their prey as they kill and devour them. natural selection!
precocial vs. altricial species
precocial: ones that can move around the environment soon after hatching, as opposed to altricial species which cannot
predator and prey sensory systems: preying mantis
predator and prey sensory systems CO-EVOLVE selection acts on each species: it favors successful predators but also favors stronger prey who can escape-->a co-evolutionary arms race example: -moths have evolved ANTIBAT SOUNDS -signals toxicity, startle bat to gain moment, jam sonar---all in different species/situations tympanal organ in insects. the organ detects sound. PREYING MANTIS HAVE THESE -analyses suggest that these types of organs evolved independently 18 different times. anti predatory response to echolocating bats -when it detects orienting cry, it either flies away or flies erratically (depending on volume, like the moths) -STUDY: put vaseline on the tympanal organ and then mantises couldn't respond to the cries at high frequencies
Alfred Russel Wallace
published before darwin did
Morgan's Canon
"... in no case may we interpret an action as the outcome of the exercise of a higher mental faculty, if it can be interpreted as the outcome of one which stands lower in the psychological scale." BUT...this by no means excludes the interpretation of a particular act as the outcome of the higher mental processes if we have evidence. not saying it's impossible...just saying we need evidence for it
Jay Gould "Just So Stories"
"Just so stories" Phrase has been used to criticize evolutionary explanations of traits that have been proposed to be adaptations but lack adequate data tempting to call a trait an adaptation without data (pink flamingo example. Pinkness comes from their diet) May have some element of truth, but crucial issue that you have data
Lloyd Morgan
"Morgan's Canon" very influential "Reductionistic approaches"! -->emergence of new field: COMPARATIVE PSYCHOLOGY Rather than grand theorizing, he argued for careful, minimalist explanations (not anthropomorphic explanations) In Animal Intelligence, Romanes said scorpion under extreme stress stinging itself to death= like human suicide. Morgan's response: this behavior is a simple reflex .... not suicide!
Darwin Bowler points
- accused by critics of concealing that he was not the first scientist to propose a theory ofevolution. - he was well aware of earlier attempts,although convinced that his own approach was superior. - Define the concept of 'evolution' -- any non-miraculous process by which new forms of life are produced - the first such theories were proposed by ancient Greeks.
Plato + Aristolal
- argued form of each species of life was absolutely fixed -for Plato each species had an ideal form or pattern whichwas conserved at a transcendental level, members merely copies of this fixed pattern. -- TYPOLOGICAL view of species, -number of absolutely fixed species - maintained their structure - could not blend by hybridization. Individualvariations were by definition trivial and transitory. --- 'chainof being' - a linear pattern stretching from the lowest formof life up to the human race.
crows & whelks
- bird dropping height minimizes total energy expenditure in upward flight - diminishing returns after 5m - but CANT quite say "crow is programmed" to chose dropping height to minimize flight (aka genes code for flight) - could be connected to learning! aka selection of brain capable of learning efficent behavior
Jarman and African ungulates
- classified 5 groups - gradient of body size, habitat, diet, group size, reproductive unit and antipredator response - all of these charactersitcs are connected!! gut size is an important component to efficent digestion of foliage - hard to digest plants tend to need larger total body size
brain size among cetaceans
- comparing brain weight relative to body weight - dolphins (almost same as humans) above regression line -baleen whales below line -connected to food gathering (hunting/vs filter feeding)
starlings at the nest (kacelnik)
- economics of foraging - how many items should parent bring on each trip to maximize rate of delivery of food to nest - MARGINAL VALUETHEOREM--- graphical solution
General History
- eighteenth century ; first comprehensive ideas of the transmutation ofspecies developed (often differed markedly from modern perspective). - early nineteenth century ; theories by Lamarck + others were beingwidely discussed by radical thinkers, -->still rejected by the conservative scientific community -
Turtles experiemnt
- in lab setting turtles move in direction based on magnetic cues for location (manipulatable) - turtles displaced upwind had much more difficult time returning to island than those placed downwind
bat responses to insect defenses
- some bats lowered frequency of sonar,making them more difficult for detection by moths - some moved to ultrahigh frequenciesabovesensivity of moths -stealth echolocation ------ aka lower intensity calls to avoid detection but there is a cost!!maximum distance of bat detection of moth decrease 3x-4x---demonstrating its solely aconteradaption to moth hearing an bears no advantage to echolocation
Crook and Weaver birds of Africa
- studied 90 species -strikiing differences (coloration, social/sexual behavior, diet, habitat) -correlational study -group 1 vs 2 - insect (solitary, monogomous, drab, forest) - seed (bright, colonies, polygamous, savannah) food abundance and distributionthought to be main driving forces
Startle hypothesis in moths
- suggest tiger moths produce deimatic display illiciting startling reflex -gain moment advantage - while bats initially startled by low-duty-cycle moth clicks, they quickly habituated -suggests startle effect is ephemeral and unliekly to be effective under natural conditions for long time however - high-duty-cycle clicks had longer lasting effects----> play important role at least for naive bats!!!
Cris sandoval and striped walking stick
- walking sticks on narrow leaf bushes developed stripe (compared to wide-leafed solid green relatives) - dorsal stripe morphology as adaptation - evolved independently 5-6 times -followed up w/ expermient measuring selection on individuals w&w/o stripes on both plants
Early Ethology
-BEHAVIOR EVOLVED. -Early goal of ethology was to demonstrate that behavior, like anatomical and physiological traits, had EVOLVED. -They stressed the ADAPTIVE nature of behavior. - Emphasis on INHERITED BEHAVIORAL response: "instinct" -interesting: Has to be at least a partial genetic basis for traits in order for evolution to be involved. INHERITED BEHAVIOR: INSTINCT Species studied=insects, birds, etc. Why so much attention to inherited behavior? -need to look at evolution -before u explore a particular behavior, you have to understand the entirety of a species -constructed Ethograms - detailed behavioral profiles. (bc you have to study the WHOLE before you can really target a particular behavior)
Richard Dawkins on Darwin
-Darwin observed artificial selection (human breeders using their eye to decide) and compared it to what goes on naturally -he saw that species don't stay the same over time -how does natural selection work? animals struggle, suffer for survival -horrors of nature (food shortage, disease, etc.) Thomas Malthus wrote about human struggle, Darwin applied those ideas to what was happening in nature. -struggle for existence: not enough resources for everyone to survive -tiniest variation could make crucial difference in animal's chances of survival-->reproduction-->pass variation down -he hesitated to publish his ideas because he knew it would be upsetting for religion
Morgans reductionist parsimony approach
romanes observed "scorpion suicide" and morgan argued he went too far- can not attribute it to level if it can be attributed to lower mental capacity
Lorenz and Tinbergen Simple Models study
-Hawk/goose experiments -simple models -Young geese duck down or run to their parents when they see hawk in the sky. Adult geese spread out their wings to form a barrier to protect young. Are these FAPs or learned? gosling responds to hawk shapeeven with no previus exposure! only respons to hawk/oriented shape (shape and direction of movement -L and T created hawk model, made it "fly" above geese. Adult geese spread wings, young ran to parents. -they discovered that it depended on which direction the hawk is moving. If moving in certain direction, the geese wouldn't respond. If moving in opposite, they would. Bc they looked like hawks when they were moving in a certain direction. conclusion: nervous system plays important role in distinguishing • Depending upon which direction the model was moving, you would either get a response form the goslings or not o Simple stimulus moving in one direction looks like a hawk, other way looks like a goose o Same model, depending upon direction of movement would either elicit or fail to elicit a response o Hawk visual plus particular movement is what elicits response o Allowed them to use two pieces of information this makes them wonder: what's up with the nervous system? the nervous system is maybe in some way defining what is crucial about the releaser (direction/type of motion). this has to do with the IRM. hirsch and cologues (comparative psych)useddomesticaeted chickens and didnt find same result (but yo they were domesticated)
Bat adaptations for echolocation (unlike typical mammal)
-Higher frequency -Has to produce a LOUD orienting cry so that it projects further into the environment. but the echo will be quieter, so you have to be loud enough that you can hear the echo o Right after the cry, the bat blocks out noise for the millisecond before it hears the echo (because its cry was so loud). this is hard because you need to be able to hear the echo, but you're blocking out sound Bc the loud cry affected your hearing -->bat adapted MUSCLES THAT SEPARATE MIDDLE EAR BONES right after the cry is produced. then it shuts it right back in time for the echo to be heard
Kruuk and Intraspecific variation in social systems
spotted hyenas - populations varied based on food availability - solitary + nomadic scavengers - social + territorial hunting groups (larger more abundent prey available) - the solitary groups form social hunting groups when prey more abundant! (ex during wildabeast migration) FACULTATIVE BEHAVIOR
facultative behavior
survival under varying enviro conditions by assuming variosu behaviors ex. spotted heyenas will go from solitary scavangers to group hunters. during wildabeast migration
Questioning Roeder's hypothesis: Fullard et al. 2003
terminal buzz: when the sound gets faster at the end (remember how it sounds) Fullard measured the buzz and end of buzz found that at the very end of the buzz, A2 does NOT fire. Roeder said that it DID fire. but their concerns are not necessarily accurate. it's possible that A2 fires and then has a recovery period. so the moth will still go into chaotic flight even though the A2 isn't firing. it doesn't have to fire. (FAP goes to completion)
Continuation of goose/hawk study on chickens
-Jerry Hirsch: "the Tinbergen hypothesis that certain specifically shaped sign stimuli innately arouse a fear response was tested on the white Leghorn chicken and found to be untenable under controlled laboratory conditions: -so who's right? Hirsch or Tinbergen? -Hirsh said that something must've been learned bc his chickens didn't respond in the same way. but...he might be wrong -these chickens are domesticated (under human care, devoid of any of the threats that might be there in the natural world). Does this mean that they lost their adaptive response bc there was no longer any need for it? But other conclusions: -No such thing as a representative species. chickens didn't act the same way as geese
phylogeny
the evolutionary development/history of taxonomic groups of organisms
European robin releasers
-Male has bright orange breast -Sets up territory in spring and excludes other males from entering that territory -How do they identify threat of other males? -2 models: 1) anatomically correct bird, but no bright color-->Does not attack. 2) tuft of cotton, bright color-->Attacks
Variation in sexual imprinting
there is a sex difference btwn males and females. Males have to learn who they should mate with. Females are born with that knowledge. -sexually dimorphic mallard ducks (non-monogamous): males of dimorphic species may rely on imprinting to identify sexual partners—but not the females. Females can't bc their fathers aren't there when they hatch. So it has to be innate for them. only the females incubate eggs, so females show innate preference but the males will prefer mates of their foster mother species -monomorphic male and female teals (monogamous)—both parents available to imprint on. so both males and females must learn--it's not innate non monogamous species=sexually dimorphic--male not present for daughters to imprint on, sons imprint on mother monogamous species=monomorphic, both parents available to imprint on example from video: adopted duck will chase after a girl who looks like his adopted mother, not him
Navigation
-Orientation over greater distances. - Process by which organism uses external cues to determine its position in reference to a destination as it moves about the environment. - Can involve both: map sense and compass sense
Stimulus filtering
unless the worm moves, ganglion cell will NOT fire Jurassic Park video--dinosaurs got blood from frog blood. dinosaur can't see you if you don't move.
Simple models
used to study releasers ethologists 1) --> Tinbergen used simple models on stickleback fish -First one: looks like stickleback but not red. Elicits no response -when there is red, almost instantaneous response -Third: doesn't look much like a fish, but is red. Almost instantaneous response 2) European robin -Male has bright orange breast -Sets up territory in spring and excludes other males from entering that territory -How do they identify threat of other males? -2 models: 1) anatomically correct bird, but no bright color-->Does not attack. 2) tuft of cotton, bright color-->Attacks 3) Lorenz and Tinbergen Greylag geese 4) red spot on herring gull bill.
bat video
we can't hear the echolocation sound the bat in the video is approaching from the wrong side because he can hear the moth through the leaf. if the moth stays still, the bat can't locate it bc it doesn't make any noise
Anthropomorphic
-Romanes's thing about ants was this -apply human qualities to animals (ex: ant is aware of other ant's pain, goes to help) -Romanes' left columns in his phylogeny (slide 44) ("mental ability" and "species") Romanes's work had both anthropomorphic stuff and anthropocentric stuff
Male Juncos and testosterone
what happens when you replace(increase) testosterone in male juncos? direct manipulation if you supplement increase testosterone level, home range expands and feeding rate decreases. so they are paying less attention to eating and more attention to marking their territory Territorial defense is increased -cost benefit analysis -more time defending=less time finding food --> cant go too far w production
Female Anolis reproduction also influenced by the social environment
-The courting display of males helps prepare the females to become sexually receptive CREWS' experiment: -Females were captured in the spring, brought into the laboratory and placed in the following housing condition: • No males • Castrated male (won't court female) • Male with dewlap removed • Intact males -Females housed with intact males came into reproductive condition faster than those in the other conditions!! additional sensitivities: -Sensitivity to season and seasonal variation -Warm versus cold spring (they can advance sexual activity if it gets warm earlier than usual) -Fall: no go, everything shuts down. -even if you artificially raise a female's estrogen level during the fall, she STILL won't do it. because her brain tells her it's not the right time
Sexual imprinting problem with whooping cranes
young whooping cranes: humans had to use hand puppets that look like their actual crane parents. So that they wont sexually imprint on humans and not be able to reproduce
FAP Facts
¥ F.A.P. (unit of inherited behavior) ¥ (generally) requires a releaser (sign stimuli)--unless it is a vacuum activity ¥ Occurrence of F.A.P. leads to a refractory period ¥ Thresholds for the response change as a function of internal conditions (example: female goose will respond to hormones-->moving around egg only at certain times of the year) ¥ Releaser-response patterns are mostly species-specific (not every animal has same releasers or FAP responses) ¥ Learning is not necessary for the occurrence of F.A.P. ¥ F.A.P.'s are primarily involved with behavior patterns that have adaptive biological functions
Thomas Belt
-a famous naturalist -ants helping other ants: an example of the type of anecdotal evidence that Romanes used -Belt's anecdotal evidence: observed ants saving another ant—said that he didn't think it was instinctual. Thought it was sympathetic help -Romanes said this observation proves fellow-feeling and sympathy
Lorenz: On Aggression
-after WW2 and holocaust, people wanted to explain aggression -Lorenz said we can learn a lot about human behavior from animals -he said that there is sometimes aggressive behavior without provocation -aggression is going to spill out in inexplicable ways sometimes--we need things like sports to let it out. -Talked about important questions in an overly simplistic way -Willing to make confident and bold statements that modern scientists wouldn't. without even doing research. controversial thinking • Based on his work with animals
acoustic aposematism
-aposmatism hypothesis suggests bats learn to associate moth clicks with chemical defense of many arctiids - naivebats must learn to associate clicks with distastefulness ; must catch/taste/drops sound producing distastefull moths and LEARN to avoid -startle responsehas opposite pattern- bats first startled but then habituate to clicks
key animals who migrate
-arctic tern="champion" of bird migration--longest route ever recorded -sea turtles -salmon -monarch butterflies
crickets
-auditory atuned to low frequency calling of mates -also atuned to highfrequency peak in sensitivity of hearing to detect/stear away from bats while searching fro mates
Annual reproductive cycle of Anolis
-during the winter (low food ), animals are sexuallyinactive/resting. ---> spring = males come out and set up/defend territories -N.S. avoid kids in food scare time -during the dormant phase of the year, saves energy to have gonads regress—they shrink and become inactive -in spring, the testes and the ovaries become active again -only reproduce when there is food available for young to survive -not reproducing all year, restricted in terms of when they will produce offspring and activity of gonads and activity of hormones corresponds to that - female receptivity stops w stimulation -courting increase receptivity (castrated vs normal vs dunlap removal) -BIOLOGICAL CLOCK component (estrogen injected fenales wont mate in fall)
Typical mammalian auditory system (not bat)
-external ear which helps channel/funnel the sound-->tympanic membrane-->chemical energy-->bones-->cochlea
Sexual imprinting (and examples)
-geese treated Lorenz as a potential mate-->he concluded that maternal imprinting and sexual imprinting were the same thing and occurred at the same time -some, but not all species show sexual imprinting example 1: Zebra finch males raised by Bengalese Greylag goose finch females will court Bengalese females. male will always prefer Bengalese finch females over zebra finch -does this mean that the males can't mate? (bc these females in other species don't want to mate w other species). No—he will "reluctantly" mate with zebra finch if Bengalese finch females aren't available example 2: Duck species where only the females incubate eggs: females show innate preference, however, the males prefer mates of their foster mother species. when both parents are avaliable in monogamous relationships, both parents are avalible to imprint on
Romanes's phylogenies (organization of them, how did he make them?)
-identify species that have achieved particular level of development -on the right, he suggests a point in human child development where these diff. things occur (compares cognitive/emotional abilities between the animals and humans of various ages) -he relied on anecdotal evidence to construct his phylogenies!!!! but only accepted facts from important people, and made sure observations were correct, and gave more weight to observations recorded more than once example: ants helping other ants (Thomas Belt!)
testes size and primate mating
-multi-male pairings = higher than predicted testes size -monogamous relationships below regression line - testes size correlates to sperm count so way of increasing offspring likelihood/fitness
Anthropocentric
-regarding humankind as the central or most important element of existence -other animals are only important bc they are related to human existence -Romanes' far right column in his phylogeny (slide 44) ("human age")
Aristotle
-scala naturae—"ladder of life" -simple organisms lower down, complexity increases as you go up -this doesn't have anything to do with evolution. but Romanes attempted to apply an evolutionary mode of thinking to this--first time anyone tried to use evolution to explain animal behavior Historical/phylogenetic approach
James Ussher in the 1650s.
-strict literal interpratation of Creationism vs john ray- sought a synthesis with religion + natural theology. The study of living things confirmed thateach species was carefully adapted to a particular way oflife, thereby displaying the Creator's wisdom and bene-volence.
Releasers (and examples)
-the environmental stimulus the prompts the organism to perform the FAP -sensory systems have evolved to act in a certain way in terms of particular situations in the environment Stimuli in environment that elicits FAPs Ethologists used simple models to study releasers 1) Male stickleback fish Tinbergen 2) European robin 3) greylag geese Lorenz and Tinbergen 4) red spot on Herring gull bill Tinbergen
Errors with Lorenz and imprinting
1) Critical period: Lorenz thought it was very narrow, within a few hours -BUT...Subsequent research shows there is a critical period, but it is much broader—a few days 2) Exposure to stimuli before hatching influences imprinting -Hatching synchrony (eggs hatching at same time) influenced as well -it makes sense for all eggs to hatch at same time so that they all have same chance to form imprinting connection. but she lays eggs at diff times so they prob won't all hatch at once -Hatching synchrony is dependent on adult femal incubation calls OR calls produced by siblings before hatching -Eggs in clutch coordinating development so they hatch at the same time -hatching synchrony study
Major methods to study hormone effects
1) Direct manipulation - Of hormone levels - Replacement of hormone following the removal of the associated gland - ex. suplmemented testosterone in juncos 2) Correlational studies - Study changes in behavior that mirror fluctuations in hormone levels - ex mouse uterus study or study of testosterone acorss birds and territorial behavior 3) Gene manipulation -remove a particular gene (maybe receptor for hormone, etc.) -Ex oxytocin study of male mice (responsible for female recognition)
Adaptations of the bat's auditory system
1) External ears differ in shapes and sizes Natural selection has shaped the faces of the bat's ears. some bats have different types of ears depending on type of echo they are going to hear. ex: fruit-eating bat has simpler ears bc fruit doesn't run away from you! - (all but very few fruit bats use echolocation) - frequency of echolocation calls varies from 8 to 215 kHz depending on bat species - vary the intensity,pulse rate, frequency-time, directionality, harmonics according to situation. this PLASTICITY allows better location/detection in diff habitats - greater specialization in habitat= greater selective pressure for defense by insects different species--fish-eating bats, meat-eating, fruit-eating, insect-eating, etc. all have different types of ears because they are looking for diff. things 2) Noses are different because of different kinds of environment they are having to navigate many species of bats produce their sounds through their noses. depends on their environment Sounds they produce have to be able to cope with different environments in which they are trying to find food
Errors of comparative psychology (listed out)
1) They wanted to create a simple linear scale of animal species from unintelligent to intelligent 2) Used "representative" and (and convenient) animals arranging them along the scale 3) Believed learning was the key to understanding animal behavior 4) Thought that general laws of behavior (that they would learn in the lab) could explain the behavior of all species -it's sometimes true, but there are directions that evolution can take that won't work based on general laws 5) Believed genetics was irrelevant to psychology 6) Believed laboratory studies were superior to field work on animal behavior - they thought that bc of rigor and control in lab studies, this was superior to anything that could be done in the field. but this isn't true 7) "The Snark was a Boojum"
Darwin's Theory of Natural Selection (1859)
1) VARIATION: Individuals within a species differ in their morphology, physiology and behavior 2) HERITABILITY: Some of this variation is heritable; on average offspring tend to resemble their parents more than other individuals in the population. 3) EXCESS OFFSPRING-->COMPETITION FOR RESOURCES AND MATES: Organisms have a huge capacity of increase in numbers; they produce far more offspring than give rise to breeding individuals. There is competition between individuals for scarce resources such as food, water, mates, etc. 4) NATURAL SELECTION As a result of this competition, some variants will leave more offspring than others. These will inherit the characteristics of their parents and so evolutionary change will take place via "natural selection." 5) ADAPTATION As a consequence of natural selection organisms will come to be adapted to their environment. Individuals that are better able to find food, water, mates, avoid predators and so on will have greater representation in future generations.
Behavioral complexity in cephalopods
1) camouflage allows cephalopod to hide from predators and to be a better predator too! CHROMATOPHORE ORGANS sense color-->change color (pigments in chromatophore cell) it mimics organisms that are a threat to predators-->predators won't come! it "mesmerizes" victims w/ colors-->can be a predator (makes them freeze) CUTTLEFISH do all of this 2) cephalopod and vertebrate eyes=similar mollusks have a lot of variation, but they are classified as mollusks (snails, octopus, etc,). they all filter-feed. they require a more effective visual system, so eyes evolve
Some major questions in evolutionary theory
1) the "species problem": how did different species evolve over time? 2) accounting for the diversity of life (ex: pinnochio frog) 3) examining changes in the molecular constitution of genes 4) explaining ADAPTIVE COMPLEXITY: complexity that is adaptive--different structures coming together in an adaptive purpose (flower example)
4 Evolutonary Theory Applications
1. Historical - oldest +least powerful approach - attempts to reconstruct behavioral phylogenies.... "behavioral homologies" Issues......- behavior doesnt fossilize, extant species ≠ ancestral, uses representative species, convergent evolution of traits 2. Evaluative Approach / testing adaptive value through experimintation - evaluate based on how it increases fitness - what must be advantage of behavior - ex. tinbergen removal of eggshells Issues......- pure speculation (ex. why are flamingos pink) 3. comparrative approach -compare related species in diff habitats OR simmilar enviro w different ancestries 4. predictaive approach - settign up hypothesis derived from evolutionary theory + testing them w/ behavioral data from extant species - think of optimality models
DAVID CREWS' THREE PATTERNS
3 ways in which reproduction can be regulated DAVID CREWS STUDIED ANOLIS LIZARD 1) The associated reproductive pattern - Characterized by a close temporal association between gonadal activity and mating e.g., rats, Anolis lizards -ex: it's adaptive for the Anolis to reproduce during spring. when not time, sexual systems shut down. they restart when spring comes 2) The dissociated reproductive pattern - Characterized by a complete temporal uncoupling of mating behavior from gonadal maturation and sex steroid hormone secretion: gametes produced after breeding season and stored for next -e.g., red-sided garter snake -video: male releases chemical that makes him seem like a female so they are distracted by him. then he mates with the females -if you castrate the male, he retains sexual interest. after a few years, he eventually loses it though 3) The constant reproductive pattern - Mature gonads are maintained for prolonged periods in a constant state of readiness (unpredictable onset of usually brief favorable breeding conditions. so you have to be ready all the time) -e.g., zebra finch (desert dwelling)
A plant that evolved a bat beckoning beacon
A rainforest vine evolved dish-shaped leaves that help attract the bats that pollinate it Tests revealed that the leaves were highly efficient at bouncing back the echolocation sound pulses of nectar feeding bats so...not only insect-eating bats use echolocation co-evolution between plant and bat
Three components of the reproductive process
ACTIVATIONAL EFFECTS OF HORMONES Three components of the reproductive process: 1) Secretion of sex steroids (hormones) by the gonads (ovaries and testes) 2) Production of gametes 3) Timing of mating behavior In some species these are tightly linked, as is the case with the ANOLIS LIZARD ... studied by David Crews
Francis Darwin
ANTHROPOMORPHIC/CENTRIC Charles Darwin's son, Francis, an eminent botanist, gave a presidential address on "consciousness in plants" to the British Botanical Association ...
Alfred Binet
ANTHROPOMORPHIC/CENTRIC The Psychic Life of Micro-Organisms (developed human intelligence testing) -interprets behavior in an anthropomorphic way -extending the argument for mental evolution -said that Micro-organisms ALSO have the same mental faculties that Romanes said animals had -Claimed that not only Romanes' animals are showing sophisticated emotions, but so are micro-organisms. even plants!
inferior colliculus
Additional adaptation for echolocation Has "echo-detector" cells that are sensitive to the sound of the echo! makes it easier to hear the quiet echo has tracking neurons
lateral lemniscus
Additional adaptation for echolocation tract of axons in the brainstem that carries information about the sound from the cochlear nucleus to various brainstem nuclei and ultimately the contralateral Neurons that block (inhibit) transmission of auditory messages to higher regions of the brain during echolocation sounds alone with middle ear bones moving, this prevents the negative impact of the loud cry sound has tracking neurons
Other details for noctuid moths
• Sound below 20 kHz. is not responded to no matter how loud. • No discrimination of different bat species frequencies in the 20-40 kHz. range • Caterpillar of noctuid moth is responsive to a 1000 Hz. sound. (completely diff than full grown moth. this is bc it is responding to the threat of a hunting wasp) UMWELT
