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Midterm #1 Study Questions - Flashcards

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Class:NPB 102 - Animal Behavior
Subject:Neurobiology,Physio & Behavior
University:University of California - Davis
Term:Fall 2012
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What were Niko Tinbergen’s “four questions” about behavior? Which of each of the four questions do the different components of the “levels of analysis” correspond to?
Proximate Causes: Mechanism
-       Genetic-developmental mechanisms: How does it develop? (Developmental)
-       Sensory-motor mechanisms: What mechanisms cause it? (Immediate/Causal)
Ultimate Causes: Evolution
-       Historical pathways leading to behavioral trait: How did it evolve? (Evolutionary history)
-       Effects of selection on history trait: How does it contribute to survival? (Fitness)
What is Tinbergen’s “Zeroth Question” and why is it important?
Tinbergen’s “Zeroth Question” is “What is the animal doing?”. This is important because in order to examine the four questions about behavior, we first have to establish that the animal is exhibiting a behavior.  The observation also leads to forming hypotheses.
What relatively unusual feature of prairie vole biology has made this species particularly interesting to study from the standpoint of the neural basis of complex behavior?
Unlike most mammalian rodent males, prairie voles are monogamous, they only mate with a single sexual partner. Prairie voles have much more numerous V1a receptors than polygamous voles. The receptor rich male voles attach themselves more strongly to females, even if they had not mated yet. (I thought the highlighted sentence refers to a special study researchers did.  This behavior is different from normal prairie voles.  Normal prairie voles form monogamous attachments from the vasopressin released in the act of mating, which binds with the V1a receptors and triggers a reward response in the brain)
What are V1a receptors, what are V1aR genes, and what significance do they each appear to play in mating behavior of prairie voles? How does the V1aR gene of male prairie and male montane voles differ, and what is thought to be the significance of this difference?
V1a receptors are vasopressin receptors in the ventral pallidum (located at the base of the brain) found in prairie-rich cells. This activity in turn affects neural pathways in the brain that provide the vole with positive feedback.
V1aR gene is present in male prairie voles, while male montane voles lack V1aR gene. The absence of this gene in montane voles can be explained as Proximate Development, for its polygamous behavior.
The textbook says that both voles have versions of the V1aR gene, but that the allele possessed by prairie voles contains an extra section.  This difference leads to higher production of v1a receptors in the brain of the male prairie voles.  Because there are more receptors, male prairie voles have an exaggerated reward response to vasopressin released after copulation with a particular female, leading to greater pair-bonding.  (correct me if I’m wrong, I may have misread) [[Anon: ^Agreed; *Note: The book doesn’t specify any difference in name for the V1aR/avpr1a gene with/without the extra DNA sequences. (Both species have avpr1a genes, but prairie voles’ V1aR have relatively more DNA)]]
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Provide one explanation for monogamy in male prairie voles at each of the 4 levels of analysis (i.e., answering each of Tinbergen’s four questions).
P-D: Throughout the development of the prairie vole from newborn to adult, an increase in the levels of vasopressin it produces can be measured, leading to an increased likelihood of monogamy.
P-C: Male prairie voles have extra DNA, specifically the avpr1a gene; the gene increases the amount of vasopressin receptors in the ventral pallidum which in turn provides positive feedback to the males when copulating with just one partner. Isnt this P-D? anyone? I thought when we talk about gene expression is proximate (gene/development)?
U-H: Prairie vole males that formed close  attachments to a mate in the past left more descendants than males with a tendency towards polygamy.  
U-F: Monogamous males have forced their mates to be faithful by guarding them, which has meant that the offspring produced almost always carry the genes of mate-guarding male.
Describe how genetic manipulations have been used to explore the importance of the V1a receptor to pair bonding behavior of both prairie and meadow voles.
Transfer of V1aR gene from prairie to montane voles, lead to increased expression of V1a receptors in ventral pallidum of montane vole brains. The genetically altered males formed much stronger attachments to a current mating partner than did unaltered control voles.
Defend or criticize the following statement: The fact that two recently-evolved species of vole, the montane and the meadow vole, are both polygynous demonstrates that the common ancestor of three species (prairie, montane and meadow vole) must have been polygynous (you’ll need to look at the figures in the textbook to answer this well).
The common ancestor is probably not polygynous. According the phylogeny tree on page 6 of textbook, the most recent common ancestor of the three species were monogamous.
What is the difference between “pure science” reasons and “applied” reasons for studying behavior (or anything else)? What are some examples of each? Why is it important to study problems from a “basic” standpoint even if your main interest is in “applied” problems?
pure science” reasons - to satisfy curiosity about something; to study behavior critical to the animal’s survival/reproductive success

(e.g. homeostasis, predator avoidance, reproduction)

-- “applied” reasons - to solve some sort of problem or use what we know to better understand something for ourselves

(e.g. understanding ourselves, control of pests/diseases, conservation/welfare)


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What is the difference between “levels or organization” and “levels of analysis?”
levels of organization include how things like molecules, organelles, cells, tissues, and organs are put together to form individuals and how individuals congregate into things like populations, communities, and ecosystems.
-- levels of analysis are different ways of answering questions about behavior; at each level, one can make multiple hypotheses that answer each of Tinbergen’s questions; each of Tinbergen’s questions can be considered to be a level of analysis (the levels of analysis include: proximate-developmental, proximate-causal, ultimate-historical, ultimate-fitness)
What was the hypothesis proposed by Tinbergen for how beewolf wasps locate their nest entrances? Propose an alternative hypothesis to this hypothesis. Now propose another hypothesis that is NOT an alternative to the first hypothesis. What distinguishes the two new hypotheses you proposed that makes one of them an alternative to the one proposed by Tinbergen and one NOT an alternative? Would Tinbergen say that the one that is not an alternative to his hypothesis is therefore not interesting? Why or why not?
Tinbergen’s hypothesis:
- To locate their hidden nest entrance, beewolf use visual landmarks for orientation. (P-C)
Alternate hypothesis: (P-C)
- They detect the nest through chemical signals produced by the dead bee inside the nest.  (Same level of analysis, but provide different possible reasons)
Non-alternative hypothesis: (P-D)
- (as the wasps grow) The wasps learned how to detect nest from observing the behavior of fellow wasps. (Different level of analysis, and 1st hypothesis could also be true)

Alternative hypothesis are at the same level of analysis as the original, and the two can contradict but they are not necessarily mutually exclusive. Non-alternative ones approach the behavior at a different level of analysis. Tinbergen would still be interested in the non-alternative hypothesis because it would provide additional information and more depth on how this behavior works.
Now go through the same exercise as for the previous example but using the hypothesis proposed by Tinbergen for why gulls and other birds remove eggshells from their nests.
Tinbergen’s Hypothesis: They remove the egg shells to reduce risk of predation on the newly hatched chick.
Alternate: (Same level of analysis U-F): They remove egg shells to prevent the baby gulls from injuring themselves.
Non-alternate: (Diff level of analysis): They remove the egg shell simply to keep the nest clean (P-C)
Following up on the egg-shell-removal example, imagine that you discovered (observed) that adults of a species of bird you’re studying eat the eggshells from their newly-hatched eggs rather than carrying them away and dropping them. Propose one new ultimate/fitness hypothesis and one new proximate/immediate causation hypothesis for “egg-shell-removal” behavior that takes into account this new information. Are either of these hypotheses “alternatives” to Tinbergen’s original hypothesis? Why or why not?
Ultimate/fitness: Eggshells contain calcium and other nutrients, so eating them will give the parents more immediate energy to take care of their young.
Proximate/immediate: The parents see eggshells in general as a viable food source.
Another proximate/immediate: The eggshells just simply taste good (sensory-motor)
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Many of you may have observed that there are huge flocks of crows that roost in the trees near the Life Sciences Building on campus (thoroughly befouling the entire area). Propose one hypothesis explaining this communal roosting behavior, and one alternative to your hypothesis. Are your hypotheses proximate or ultimate? Which of Tinbergen’s questions do they answer?
Hypothesis: The crows roost together so that they can stay warmer overnight.
Alternative:  The crows roost together to confuse predators by making it harder to focus on only one bird.
These are both ultimate-fitness hypothesis, and answer “how does it contribute to survival?”
What is the syrinx, and how is it used by birds to produce sounds? Be as specific as you can about how different parts contribute to sound production.
The syrinx is the primary sound-producing organ in birds, located at the junction between the trachea and the two bronchi. It is complex, containing many separate muscles, the most important of which (to us, at least) push a piece of cartilage in either bronchi to pinch it off. Sound is produced when one bronchi is only partially pinched and air is forced through, causing the medial and lateral labium to vibrate. High frequency sounds are produced when the left bronchus is completely pinched and the right one is only partially pinched. Low frequency sounds are produced when the right bronchus is completely pinched and the left one is only partially pinched. The phenomena of two voices occurs when both right and left medial labia are only partially pinched off so that a different note is coming out from each side. The “two voices” are simultaneous double tones that are not harmonically related and therefore must be derived from two independent acoustic sources.
Imagine an experiment where you begin with a fully adult male bird that has crystallized its song, and you then infuse a local anesthetic into the syrinx that specifically targets the muscles controlling the left side. This procedure numbs those muscles and prevents the bird from regulating air flow through the left bronchus. That is, the left bronchus would remain completely open all the time because the bird cannot contract the left syringeal muscles. How do you think this procedure would affect the song of the bird? Be as specific as you can about the effects it would have on the different elements of the song. What would happen if instead you inserted a little plug into the left bronchus to prevent all air flow through it?
At the review session, he said that no sound would be made, since the left bronchus is fully open.
Explain what is meant by the statement that songbirds have “two voices” and describe anatomical and experimental evidence consistent with this idea.
Sound is produced when one bronchus is only partially pinched and causes the medial and lateral labium to vibrate. High frequency sounds are produced when the left bronchus is completely pinched and the right one is only partially pinched (only the right set of labia vibrate). Low frequency sounds are produced when the right bronchus is completely pinched and the left one is only partially pinched (only the left vibrate). The two sides are under independent control, and a wide variety of pitches can be made by manipulating how pinched either bronchi are. The phenomena of two voices occurs when both right and left medial labia are only partially pinched off so that a different pitch is coming out from each side. The “two voices” sound like simultaneous double tones that are not harmonically related and therefore must be derived from two independent acoustic sources.
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Use the example of red-winged blackbird song to explain why Tinbergen’s choice of the term “survival value” was not ideal. What term would you suggest using instead, and why?
The female bird has to distinguish between real redwing blackbird calls and fake mocking bird calls this increased fitness more so than survival value. The female will still survive if she mates with the wrong species, but her fitness in producing viable offspring goes way down.

The female bird right?
The differences between song types has more to do with sexual selection “ornaments” than natural selection.

Describe how the song type matching studies of free-living song sparrows clearly have both proximate and ultimate interpretations.
I think what he is asking is that in the past, males have differed in their song control systems and thus in their behavior. Some of these differences have become hereditary and some males have hereditarily been better than others at responding aggressively to save their territory better or their capacity to cope with rival males. These males’ superior behavioral skills have translated into greater genetic contributions to the next generation, where those genes have been available to participate in the developmental process within members of that generation. Therefore the hereditary attributes of those animals will in turn be measured against one another today in terms of their ability to promote genetic success in the current selection round. This is how it has both proximate and ultimate interpretations.
(similar answer in the book on page 58)
This is speculation: ultimate- being able to successfully type match is how a bird can aggressively respond and try to save it’s territory from other birds.
“Open Ended Song Learners” are believed to be able to keep memorizing new songs well beyond their first year of life, and possibly throughout their lives. One of your friends tells you that song sparrows must be “open ended song learners” and cites the evidence that young males establishing a territory for the first time appear to be able to crystallize songs that are close matches to those of their neighbor males. How would you respond to this assertion?
I would say that the sparrow most likely socialized with other species in the territory since socialization influences the behavior of the bird (page 49-52). Their songs crystallized when they’re establishing their territory for the first time, but they’re still in the end of the sensory-motor phase and are still overproducing and end up narrowing down their song choices based on their neighbors song.

I believe Hahn described this in OH as they learn songs 1-8 in critical period. Then when they are establishing territory and their neighbors are singing 3 5 and 7 it will crystallize those songs. I would say that the birds needed to have been exposed to those songs during critical period, to be able to crystallize. As far as different dialect?? any ideas beyond each neighbor has a different dialect?  

I think that the dialect evolved based on the environment.  The bird will begin to produce songs based on the acoustics of the environment.  An example is that birds living in the city will produce fast, high pitched songs whereas birds in the country or forest will produce longer, lower frequency songs.

Define a song dialect, and contrast song dialects with at least two other forms of inter-individual variation in song. Explain what kind of information you would need in order to distinguish among all of these forms of interindividual variation in song.
Song Dialect-Different song types, may be “marked off” in different geological areas. Must occur with distinct markers --> 1. Variation within an area less than variation between areas. 2. Boundaries between song variants abrupt.
The answer key says “A song dialect is a form of inter-individual variation.”
Inter-individual variation (Non-geographic Variation)-song varies only on individual basis, no distinct areas of same song types. Multiple individuals in one area could have multiple different song types.
Clinal Variation (Non-dialectal Geographic variation)- song type gradually changes as you move through an area. Variation in song types in areas next to each other are very difficult to distinguish, but variation in song types of areas at either end of cline more easily distinguished. (further apart, more variation)

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Give one proximate and one ultimate explanation for why the songs of birds living in different habitats (e.g., densely-vegetated versus more open) might emphasize different sound pitches. Do the same for birds living in urban versus wild woodland habitats.
Vegetated vs. open:
-Proximate- Growing up, bird listens to ,say high frequency song and therefore, learn to sing high frequency
-Ultimate-Densely vegetated areas have more barriers to sound, lower frequency sound is transmitted more effectively. Over time, the birds have adapted to using lower frequency for this reason.
Urban vs wild:
-Proximate- Young birds are likely to acquire the song types they can most easily hear, and so urban birds hear high-pitched songs, therefore develop a song in the high-frequency range.
-Ultimate-  Urban environments are filled with sounds in the lower-pitched frequencies.  This is why urban birds use higher-pitched vocals so they can be better heard by potential mates or enemies.
What are the stages of song learning in male songbirds, and what is going on in each stage?
There are three stages of song learning in male songbirds:
-The first is the sensory phase where the bird must hear model songs to copy during the critical period. The bird will have a strong preference for conspecific song and is very sensitive to the sequence of phrase pairing of models.
-The second stage is the sensory-motor phase where the bird must practice producing songs and be able to hear to match tutor songs. In this phase, the bird over-produces its song (produces a greater range of what they sing).
-The last stage is the crystallization phase where the bird selects song(s) to perfect and sing. Social environment affects song selection from the overproduced repertoire.
One of the key features of so-called “critical period” songbirds is that memorization of new songs is only supposed to be possible during a brief “critical period” early in life. Imagine that you are studying a species of songbird believed to be a critical period learner, but that you discover that the song repertoire of males tends to increase from one year to the next as the males age. Could this increase in song repertoire with age occur even if the birds really are critical period learners? If not, why not? And if so, how?
Many wild songbirds hear many different songs during their critical period. During their initial crystallization stage, they choose only a few songs to crystalize. However, their brains become more plastic over winter, and then when they get back into the spring season and get new neighbors, if one of their neighbors are singing a song they haven’t crystallized but they had heard during their critical period as a baby, they may decide to bring up that memory and crystalize that song. This means that they have added a song to their repertoire! However, they didn’t learn the song they heard as an adult, the song must have been brought up from memory and crystallized.
What is a “neural template” for song learning? Describe how you think the “neural template” might function in song learning of an individual male bird that has received tutoring by tape recordings of conspecific males as compared with in an individual male that has received no tutoring at all. A good answer will discuss the hypothesized role of the “neural template” in both motor production and learning.
The neural template is the guiding principle in the nervous system that is responsible for the motor output of the song and how it’s learned. it guides the process of learning and production. In different birds it produces different functions. In the learners it acts as more of a guide during the learning process of songs, but in suboscines like the eastern phoebe it is the entire blueprint of the song because they are non-learners. ....idk about the role of motor production.....
During the sensory-motor phase, the birds refer to this neural template (along with the songs they learned by tutoring if they received it) and correct themselves based off it.
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Which of the following statements would you say better describes the “neural template” for (a) a white-crowned sparrow (an oscine songbird) and (b) an eastern phoebe (a sub-oscine songbird)? 1. The neural template provides a blueprint for what the bird is going to sing. 2. The neural template provides a guide for the learning process. Defend your answer.
(a) A white-crowned sparrow - 2. The neural template provides a guide for the learning process.  → White-crowned sparrows are actual song learners.
(b) An eastern phoebe - 1. The neural template provides a blueprint for what the bird is going to sing. → Eastern phoebes are vocal non-learners.
Describe some evidence suggesting that female songbirds may use male song characteristics to evaluate the “quality” of the singer.
(Not sure if this is the experiment hes referring to) Tested female attraction to males that were tape tutored. Tested with males that produced good copies of the tape tutor song and males that produced poor copies. Females (wild) were shown to favor males that produced good copies of the tape tutored song.
Describe the evidence that male white-crowned sparrows do not need to be tutored with complete songs in order to learn to reproduce essentially normal complete songs. What does this tell us about the nature of the “template” for song learning/production and how it influences the nature of songs produced during adulthood?
The slide called “Syntax Assembly from Partial Song Models” chapter 2A. Their neural template is more of a guideline (compared to the phoebe’s full on map). Although only fragment of a song was played during the critical period, the wcs used both its innate and auditory cues they hear from the song to figure out what they should sound like.  
Describe the evidence that female red-winged blackbirds are more discriminating when it comes to song than males are. Provide one proximate and one ultimate explanation for this fact.
Expt.1 Female red-wing blackbirds tested for attraction displays to song of male red-winged blackbird vs. a mockingbird imitation. Found that females were able to distinguish between the real song and mimicked song-showed more displays to male RWB’s than mockingbirds.
Expt 2. measured male RWB responses to fellow male RWB songs or to mockingbird imitation. Showed that males were less able to distinguish between the two types.
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What is meant by the phrase “interactive theory of development?” Explain how this theory involves both the “internal environment” and the “external environment.”
The interactive theory of development states that phenotype is a product of the interactions between genotype and the environment. “Environment” includes both internal (chemical environment in the body, for example hormones, neurotransmitters, and nutrients) and external (outside the body) components.
Describe evidence suggesting that changes in worker bee behavioral phenotype are related to changes in gene activation. Explain how this example illustrates the “interactive theory of development.”
The bee’s genotype responds to the environment in ways that influences the development of the bee’s phenotype (which includes proximate mechanisms underlying behavior, such as the nervous system and behavioral traits). Within three weeks, the bee’s phenotype changes from cleaning cells to feeding larvae and nestmates to packing pollen and lastly, to foraging. Studies by Whitfield have shown that comparison of the genes at various phenotypic changes yield changes in genetic activity. This example illustrates the “interactive theory of development” because it shows how the bee’s phenotype is a direct result of its interactions between environment and genotype. If you place a bee in an environment composed of young bees, that bee will turn off the genes for cleaning and feeding faster than normal and start expressing the genes for foraging at an earlier age. Conversely if there are higher than normal proportion of older bees then the bee will clean and feed for a longer period of time (express the genes for younger bee activity for a longer period of time).
Describe three hypotheses to explain the presence of blackcap warblers (which historically always wintered in sub-Saharan Africa) in Britain during the winter. Which of these hypotheses was favored by the researchers who studied this phenomenon, and how have those researchers managed to narrow in on that explanation? What assumption involving the rules the birds follow for choosing orientation direction in the autumn must be true for the researcher to be able to draw the conclusion they did from the experiment they did? Propose a straightforward experiment that would test for the validity of this assumption, and describe specifically how you would interpret different possible outcomes of this experiment. Hint: You need to deal with the concept of navigation to answer this well.

1. The birds lost migratory abilities (and just stayed in UK year round) → this was ruled out

2. Birds were migrating from Scandinavia- Travel southwest, end migration in UK

3. Birds traveled from Central Europe- would mean that birds were orienting north-west.

 → Expt.) took adult birds wintering in britain. Found they oriented northwest. When breeded, found that offspring also oriented northwest.

 --if birds were from scandinavia, they should have oriented south-west. So this supports hypothesis 3, that the birds came from central europe.

 --also defends assumption that the birds don’t “navigate”. Orient toward a direction, not destination.


Given that migratory behavior of blackcap warblers has a genetic basis, how would a new population wintering in a new area (e.g., Britain) originate? A good answer to this question will describe a logical evolutionary scenario involving the adaptive significance of migratory phenotypic traits and how it affects evolutionary change in the population.


Genetic variation exists in every population, and has effect on natural selection. (Other genetic effects on natural selection include genetic drift, bottlenecking, etc, but these are probably not playing a role here.) Basically, there are always a few birds who don’t quite follow along with the average thing everyone else is doing. Usually these birds (who migrate to different places) just die and don’t pass on their genes. But every once in a while, they migrate to a place where they can survive and pass on their genes. Consider this a type of directional selection, favoring birds that would potentially respond well to climate change with their flexible genotypic plasticity.
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A graph of “mid parent residual onset of migratory activity” versus “mid offspring residual onset of migratory activity” reveals a positive relationship with substantial scatter around the best-fit line. What is the primary significance of the positive relationship? What do you think is significant about the substantial scatter around the best fit line?

Positive relationship- suggests that genetics play a large role in bird migratory activity. Ex.) Parents with early onset migratory activity tend to have offspring that show early onset migratory activity.

Scatter around best fit line-Suggests that the environment is also a factor. If genetics were the only basis, then there would essentially be no scatter, or at least much less.

Graph also implies that the genotype of the parent does not exactly predict the genotype of the offspring. There is still genetic variation in a population.
Describe evidence indicating that timing of onset of migratory behavior in blackcap warblers is responsive to selection. How could these experiments have been made even more convincing? Describe another example of an experimental study, using a more tractable study system than the blackcap warblers, that provides much more convincing evidence that genetically heritable behavioral traits of populations can be modified by selection.

Experiment-->used directional selection to select for late onset migratory activity. Graph shows three generations-->each generation shows a later onset.

-therefore: directional selection for later migration onset produces generations of later migrants. Furthering evidence of migratory onset being a genetically heritable behavioral trait.

The mouse study on nest size was more “tractable” because it included a control, many more generations, and multiple trials/ repetitions, which demonstrated the reliability of the results. (The results weren’t due to random chance.)


Consider the mouse nest-building selection study described in lecture and in the text. Imagine that the results were obtained with all of the mice held on a constant temperature of 20oC. Now imagine that after the 15 generations of selection shown in the figure that a number of individuals from the “high” and “low” selected lineages are tested for nest building behavior at a range of temperatures from 5oC to 35oC. Imagine that the results show that mice from the “low” selected line build bigger nests at 5oC than they do at 35oC, the mice of the “high” selected line build the same size nests irrespective of what temperature they’re held on, and the two lineages build identical sized nests at 5oC. Discuss the implications of these hypothetical results and specifically address the question of whether these results are consistent with the hypothesis that nest building behavior has a genetic basis. Draw and label a figure illustrating the described hypothetical relationship between nest size, environmental temperature and genetic line.
Contradicts the evidence of the nest building having a genetic basis. Suggests that, in some cases, environment also plays a role...
He drew this out during office hours it was like the lower nest one was variable based on temperature - the colder it got the higher they went, but the high nest mice built the same nest regardless of temperature. So OP is right this is showing that environment plays a role. Environment plays a role in both the low nests (via temp) and the high nests (via nutrition for example)
The same mice species and original population, randomly split up and submitted to different directional selection treatments, where after selection they behave differently to same temperature stimulus. This implies there must have been allelic variation in multiple genes making up the nest-building behavior, in the original population. The “high” lineage mice always built the same size nest, which implies the behavior has a stronger genetic influence, like the mice are fulfilling a pre-set genetic program, and have an inflexible genotype. The “low” lineage mice are building larger nests at smaller temperatures and vice-versa, which implies the behavior has a strong environmental influence. Even within the (probably many) genes coding for this behavior, some allelic gene variants are responsive to change, and some allelic gene variants are completely inflexible to change (in environmental conditions).... I think this is a basic theory of behavior but I can’t remember what it’s called........

After the directional selection, the “high” and “low” populations could be considered different genotypes. In that case, these results imply that each population experiences a different “reaction norm,” where some genotypes are very resistant and other genotypes produce a large range of behaviors depending on environmental input.
Draw a graph illustrating what you think would happen to the mean value and the variability in nest-building behavior in mice following a number of generations of directional selection on the trait. Do the same if the selection was stabilizing.

Variability would increase from the directional selection treatment.

Variability would decrease from stabilizing selection treatment (mean value would even out and be representative of the whole group).
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Which of the following two statements regarding the control of movement behavior of fruit fly larvae is more reasonable, and why? (1) Wandering behavior is controlled by a single gene locus. (2) Inter- individual differences in wandering behavior can be accounted for by differences at a single gene locus. Explain your choice.
Behavior is best explained by an interaction of genes and the environment. However, inter-individual DIFFERENCES can be attributed to just genetics. Therefore, option 2 would be the best choice. Variances in such behavior also attests to this.
How do coast and inland garter snakes differ in their interest in, and willingness to eat, banana slugs? Consider the hypothesis that experience with slug-derived chemicals in utero accounts for the behavioral differences observed between coast and inland garter snakes. Propose an alternative hypothesis, and make one prediction for each hypothesis that would help you decide which hypothesis is the best explanation.
Coast garter-Love and prefer to eat banana slugs
Inland garter-large aversion to/avoid eating banana slugs
Alternative (must be proximate-developmental)--baby coast snakes learn to eat slugs from their mothers (after birth, not in utero).
Prediction for original-baby coast snakes would already have a preference for slugs, right after birth.
OR, make sure pregnant mothers do not eat any banana slugs. If hypothesis true, these babies would not show a preference for slugs after birth
Prediction for alternative-baby coast snakes would not show a preference for slugs if taken away from mothers right after birth. Would only show preference after “shown” by mothers to eat banana slugs.
Contrast organizational and activational effects of hormones and give one example of each.
Organizational effects: contribute to establishment of neuro-muscular and morphological aspects, important for behaviors to be expressed
- Usually occur during a critical period (ie, a period when if hormone not present, may not be responsive to it in future)
- Defined as irreversible (like critical period song learning in birds-- always stuck with the effects)
- Tend to be big morphological effects (degree of early exposure to hormones determine whether a structure is present at all, or functional)

Activational effects can happen any time, not just as adults; opposite of organizational
- Immediate causal, short term effects
- Effects not permanent, reversible, no critical period
- purpose is to modulate activity of existing machinery and how it functions (example of testosterone effecting plasticity of bird song)
Describe how the embryos in a pregnant rat might influence the development of one another through hormones. Would these effects be organizational or activational? What specifically are 0M, 1M and 2M embryos? What role does alpha fetoprotein play in this system?
Babies “soaking” in maternal estradiol. Estradiol normally masculinizes, which you don’t want because need some females-->babies produce alpha fetoprotein to bind and inactivate estradiol
- But does not bind testosterone--> in brain or other target tissues testosterone is locally converted into estradiol and does its business
- Male testosterone goes into the blood surrounding the embryo, and can go into nearby female and affect their brain too
 -0M=2 Female surrounded by 2 other females (0 males)
 -1M=Female surrounded by 1 male and 1 female
 -2M=Female surrounded by 2 males
   -->2M female more masculinized because brains exposed to more testosterone than 1M or 0M females (actually the estrogen thats having the effect, testosterone is the precursor to estrogen)
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What is lordosis? How have studies using lordosis and aggression as behavioral assays been used to evaluate organizational effects of hormones?
Lordosis - when fertile, female rats arch back and deflects tail to side to facilitate copulation with male.
Cite experimental data to evaluate the following statement: Laboratory rats can be conditioned to perform any naturally-occurring behavior to avoid an unpleasant stimulus, such as a foot shock.
From the notes: “Rats can be readily trained to run or turn in order to avoid a mild electrical shock. Both running and turning are appropriate natural behaviors for rats to display to avoid things like predators or other frightening stimuli. IN CONTRAST, they appear unable to learn to rear up on their hind legs- a behavior naturally used during exploration for food, not for escape from frightening stimuli. This illustrates that certain types of association are much more easily learned than others.”
---> So, this would be false. They can’t be conditioned to perform ANY naturally occurring behavior. Only those behaviors that would normally be associated with an unpleasant stimulus (running or turning).
. One of your friends says, “I think the difference between Clark’s nutcrackers and other related birds like pinyon jays and scrub jays in ability to learn tasks requiring spatial memory just reflects the fact that Clark’s nutcrackers are generally better learners than those other species, not that there’s anything special about their ability to perform spatial memory tasks specifically.” How would you respond to this?
Studies were done on Nutcrackers, pinyon jays, and scrub jays to test both their spatial memory abilities and non-spatial memory abilities. For the spatial memory task, the nutcracker significantly outperformed the other birds, suggesting a superior spatial memory. On the non-spatial memory task, nutcrackers did not exceed the abilities of the other birds. Comparing the two tests together suggests that the nutcrackers have a specialized spatial memory. If they were “simply better at learning” then they would have outperformed the other birds in both the spatial, and non-spatial memory tasks...but they didn’t.
In response to your answer to the above question, another one of your friends says, “Well, I think the differences between Clark’s nutcrackers and other jays are really primarily environmentally determined. Nutcrackers are just motivated to do a lot more food caching when they’re young than the other jays are, so they get more experience caching and retrieving caches, so they get better at it.” How would you respond to this? Propose an experiment that would allow you to determine whether or not this concern is justified.
You could capture some baby nutcrackers, before they have had the ability to “experience caching and retrieving”. Test their spatial abilities in the lab. If their spatial memory still seems to be significantly superior to other birds, than this could be attributed to more genetic factors than environmental
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What is the “nutritional stress hypothesis” with respect to song learning in songbirds? Explain how this hypothesis has both proximate and ultimate components.
One thing that could constrain song characteristics to be honest indicators of male quality relates to the fact that specific brain areas are critical to the learning and production of song, and owing to the extremely rapid development of these brain areas (young songbirds have exceptionally rapid growth and development) there is terrific potential for their functionality to be compromised if young males are exposed to any kind of  nutritional deficit at critical development stages. Thus, only males that were well cared for during the important period of very rapid growth and development and/or males that are particularly resistant to disruptive effects of nutritional deficits during that time – i.e., males likely to be “high quality” individuals – would be able to develop brain regions maximally capable of learning and producing songs. In particular, features of song such as how well tutor songs can be copied, or the size of the repertoire of songs that can be remembered, could be negatively affected by exposure to nutritional stress during development.”
Ultimate component- Females will only choose males with a “higher quality” song. Stress during development could affect the quality of song produced by males and negatively affect their reproductive success with females.
Proximate component- Nutrition during development immediately correlates with the quality of song produced by males later on.
What is the relationship between feather growth and first year song repertoire in great reed warblers, how was this relationship demonstrated, and how does this finding either support or refute the nutritional stress hypothesis regarding song learning in birds?
The longer the feather length when younger, shows that the bird is either resistant to poor nutrition or getting a lot of nutrition. When the birds are all grown up those with the longest feather length also have the most popular song, this supports the nutritional stress hypothesis because the birds with shorter feathers also had poorer songs.
Propose an experimental study that could test the validity of the nutritional stress hypothesis (Hint:Look at Figure 2.29 on page 56 for an idea.)
Count the number of female precopulatory displays compared for a group well fed/highly resistant to nutritional stress birds versus. birds that were not given good nutrition during development??
Beyond providing a potential explanation for variation in the abilities of individual male songbirds to copy and produce tutor songs, what is the ultimate importance of the nutritional stress hypothesis? (Hint: It has to do with the question of why female birds rely at least partly on song for choosing mates.)
because it is an honest indicator of the male bird’s overall health
maybe-ULTIMATE importance--> affects reproductive success with females. Acts as an indicator to females of the health of males. Want to choose healthy males for better fitness of offspring.
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 What were Niko Tinbergen’s “four questions” about behavior? Which of each of the four questions do the different components of the “levels of analysis” correspond to?
Proximate Causes: Mechanism
-       Genetic-developmental mechanisms: How does it develop? (Developmental)
-       Sensory-motor mechanisms: What mechanisms cause it? (Immediate/Causal)
Ultimate Causes: Evolution
-       Historical pathways leading to behavioral trait: How did it evolve? (Evolutionary history)
-       Effects of selection on history trait: How does it contribute to survival? (Fitness)
 What is Tinbergen’s “Zeroth Question” and why is it important?
Tinbergen’s “Zeroth Question” is “What is the animal doing?”. This is important because in order to examine the four questions about behavior, we first have to establish that the animal is exhibiting a behavior.  The observation also leads to forming hypotheses.
 What relatively unusual feature of prairie vole biology has made this species particularly interesting to study from the standpoint of the neural basis of complex behavior?
Unlike most mammalian rodent males, prairie voles are monogamous, they only mate with a single sexual partner. Prairie voles have much more numerous V1a receptors than polygamous voles. The receptor rich male voles attach themselves more strongly to females, even if they had not mated yet. (I thought the highlighted sentence refers to a special study researchers did.  This behavior is different from normal prairie voles.  Normal prairie voles form monogamous attachments from the vasopressin released in the act of mating, which binds with the V1a receptors and triggers a reward response in the brain)
 What are V1a receptors, what are V1aR genes, and what significance do they each appear to play in mating behavior of prairie voles? How does the V1aR gene of male prairie and male montane voles differ, and what is thought to be the significance of this difference?
V1a receptors are vasopressin receptors in the ventral pallidum (located at the base of the brain) found in prairie-rich cells. This activity in turn affects neural pathways in the brain that provide the vole with positive feedback.
V1aR gene is present in male prairie voles, while male montane voles lack V1aR gene. The absence of this gene in montane voles can be explained as Proximate Development, for its polygamous behavior.
The textbook says that both voles have versions of the V1aR gene, but that the allele possessed by prairie voles contains an extra section.  This difference leads to higher production of v1a receptors in the brain of the male prairie voles.  Because there are more receptors, male prairie voles have an exaggerated reward response to vasopressin released after copulation with a particular female, leading to greater pair-bonding.  (correct me if I’m wrong, I may have misread) [[Anon: ^Agreed; *Note: The book doesn’t specify any difference in name for the V1aR/avpr1a gene with/without the extra DNA sequences. (Both species have avpr1a genes, but prairie voles’ V1aR have relatively more DNA)]]
 Provide one explanation for monogamy in male prairie voles at each of the 4 levels of analysis (i.e., answering each of Tinbergen’s four questions).
P-D: Throughout the development of the prairie vole from newborn to adult, an increase in the levels of vasopressin it produces can be measured, leading to an increased likelihood of monogamy.
P-C: Male prairie voles have extra DNA, specifically the avpr1a gene; the gene increases the amount of vasopressin receptors in the ventral pallidum which in turn provides positive feedback to the males when copulating with just one partner. Isnt this P-D? anyone? I thought when we talk about gene expression is proximate (gene/development)?
U-H: Prairie vole males that formed close  attachments to a mate in the past left more descendants than males with a tendency towards polygamy.  
U-F: Monogamous males have forced their mates to be faithful by guarding them, which has meant that the offspring produced almost always carry the genes of mate-guarding male.
 Describe how genetic manipulations have been used to explore the importance of the V1a receptor to pair bonding behavior of both prairie and meadow voles.
Transfer of V1aR gene from prairie to montane voles, lead to increased expression of V1a receptors in ventral pallidum of montane vole brains. The genetically altered males formed much stronger attachments to a current mating partner than did unaltered control voles.
 Defend or criticize the following statement: The fact that two recently-evolved species of vole, the montane and the meadow vole, are both polygynous demonstrates that the common ancestor of three species (prairie, montane and meadow vole) must have been polygynous (you’ll need to look at the figures in the textbook to answer this well).
The common ancestor is probably not polygynous. According the phylogeny tree on page 6 of textbook, the most recent common ancestor of the three species were monogamous.
 What is the difference between “pure science” reasons and “applied” reasons for studying behavior (or anything else)? What are some examples of each? Why is it important to study problems from a “basic” standpoint even if your main interest is in “applied” problems?
pure science” reasons - to satisfy curiosity about something; to study behavior critical to the animal’s survival/reproductive success

(e.g. homeostasis, predator avoidance, reproduction)

-- “applied” reasons - to solve some sort of problem or use what we know to better understand something for ourselves

(e.g. understanding ourselves, control of pests/diseases, conservation/welfare)


 What is the difference between “levels or organization” and “levels of analysis?”
levels of organization include how things like molecules, organelles, cells, tissues, and organs are put together to form individuals and how individuals congregate into things like populations, communities, and ecosystems.
-- levels of analysis are different ways of answering questions about behavior; at each level, one can make multiple hypotheses that answer each of Tinbergen’s questions; each of Tinbergen’s questions can be considered to be a level of analysis (the levels of analysis include: proximate-developmental, proximate-causal, ultimate-historical, ultimate-fitness)
 What was the hypothesis proposed by Tinbergen for how beewolf wasps locate their nest entrances? Propose an alternative hypothesis to this hypothesis. Now propose another hypothesis that is NOT an alternative to the first hypothesis. What distinguishes the two new hypotheses you proposed that makes one of them an alternative to the one proposed by Tinbergen and one NOT an alternative? Would Tinbergen say that the one that is not an alternative to his hypothesis is therefore not interesting? Why or why not?
Tinbergen’s hypothesis:
- To locate their hidden nest entrance, beewolf use visual landmarks for orientation. (P-C)
Alternate hypothesis: (P-C)
- They detect the nest through chemical signals produced by the dead bee inside the nest.  (Same level of analysis, but provide different possible reasons)
Non-alternative hypothesis: (P-D)
- (as the wasps grow) The wasps learned how to detect nest from observing the behavior of fellow wasps. (Different level of analysis, and 1st hypothesis could also be true)

Alternative hypothesis are at the same level of analysis as the original, and the two can contradict but they are not necessarily mutually exclusive. Non-alternative ones approach the behavior at a different level of analysis. Tinbergen would still be interested in the non-alternative hypothesis because it would provide additional information and more depth on how this behavior works.
 Now go through the same exercise as for the previous example but using the hypothesis proposed by Tinbergen for why gulls and other birds remove eggshells from their nests.
Tinbergen’s Hypothesis: They remove the egg shells to reduce risk of predation on the newly hatched chick.
Alternate: (Same level of analysis U-F): They remove egg shells to prevent the baby gulls from injuring themselves.
Non-alternate: (Diff level of analysis): They remove the egg shell simply to keep the nest clean (P-C)
 Following up on the egg-shell-removal example, imagine that you discovered (observed) that adults of a species of bird you’re studying eat the eggshells from their newly-hatched eggs rather than carrying them away and dropping them. Propose one new ultimate/fitness hypothesis and one new proximate/immediate causation hypothesis for “egg-shell-removal” behavior that takes into account this new information. Are either of these hypotheses “alternatives” to Tinbergen’s original hypothesis? Why or why not?
Ultimate/fitness: Eggshells contain calcium and other nutrients, so eating them will give the parents more immediate energy to take care of their young.
Proximate/immediate: The parents see eggshells in general as a viable food source.
Another proximate/immediate: The eggshells just simply taste good (sensory-motor)
 Many of you may have observed that there are huge flocks of crows that roost in the trees near the Life Sciences Building on campus (thoroughly befouling the entire area). Propose one hypothesis explaining this communal roosting behavior, and one alternative to your hypothesis. Are your hypotheses proximate or ultimate? Which of Tinbergen’s questions do they answer?
Hypothesis: The crows roost together so that they can stay warmer overnight.
Alternative:  The crows roost together to confuse predators by making it harder to focus on only one bird.
These are both ultimate-fitness hypothesis, and answer “how does it contribute to survival?”
 What is the syrinx, and how is it used by birds to produce sounds? Be as specific as you can about how different parts contribute to sound production.
The syrinx is the primary sound-producing organ in birds, located at the junction between the trachea and the two bronchi. It is complex, containing many separate muscles, the most important of which (to us, at least) push a piece of cartilage in either bronchi to pinch it off. Sound is produced when one bronchi is only partially pinched and air is forced through, causing the medial and lateral labium to vibrate. High frequency sounds are produced when the left bronchus is completely pinched and the right one is only partially pinched. Low frequency sounds are produced when the right bronchus is completely pinched and the left one is only partially pinched. The phenomena of two voices occurs when both right and left medial labia are only partially pinched off so that a different note is coming out from each side. The “two voices” are simultaneous double tones that are not harmonically related and therefore must be derived from two independent acoustic sources.
 Imagine an experiment where you begin with a fully adult male bird that has crystallized its song, and you then infuse a local anesthetic into the syrinx that specifically targets the muscles controlling the left side. This procedure numbs those muscles and prevents the bird from regulating air flow through the left bronchus. That is, the left bronchus would remain completely open all the time because the bird cannot contract the left syringeal muscles. How do you think this procedure would affect the song of the bird? Be as specific as you can about the effects it would have on the different elements of the song. What would happen if instead you inserted a little plug into the left bronchus to prevent all air flow through it?
At the review session, he said that no sound would be made, since the left bronchus is fully open.
 Explain what is meant by the statement that songbirds have “two voices” and describe anatomical and experimental evidence consistent with this idea.
Sound is produced when one bronchus is only partially pinched and causes the medial and lateral labium to vibrate. High frequency sounds are produced when the left bronchus is completely pinched and the right one is only partially pinched (only the right set of labia vibrate). Low frequency sounds are produced when the right bronchus is completely pinched and the left one is only partially pinched (only the left vibrate). The two sides are under independent control, and a wide variety of pitches can be made by manipulating how pinched either bronchi are. The phenomena of two voices occurs when both right and left medial labia are only partially pinched off so that a different pitch is coming out from each side. The “two voices” sound like simultaneous double tones that are not harmonically related and therefore must be derived from two independent acoustic sources.
 Use the example of red-winged blackbird song to explain why Tinbergen’s choice of the term “survival value” was not ideal. What term would you suggest using instead, and why?
The female bird has to distinguish between real redwing blackbird calls and fake mocking bird calls this increased fitness more so than survival value. The female will still survive if she mates with the wrong species, but her fitness in producing viable offspring goes way down.

The female bird right?
The differences between song types has more to do with sexual selection “ornaments” than natural selection.

 Describe how the song type matching studies of free-living song sparrows clearly have both proximate and ultimate interpretations.
I think what he is asking is that in the past, males have differed in their song control systems and thus in their behavior. Some of these differences have become hereditary and some males have hereditarily been better than others at responding aggressively to save their territory better or their capacity to cope with rival males. These males’ superior behavioral skills have translated into greater genetic contributions to the next generation, where those genes have been available to participate in the developmental process within members of that generation. Therefore the hereditary attributes of those animals will in turn be measured against one another today in terms of their ability to promote genetic success in the current selection round. This is how it has both proximate and ultimate interpretations.
(similar answer in the book on page 58)
This is speculation: ultimate- being able to successfully type match is how a bird can aggressively respond and try to save it’s territory from other birds.
 “Open Ended Song Learners” are believed to be able to keep memorizing new songs well beyond their first year of life, and possibly throughout their lives. One of your friends tells you that song sparrows must be “open ended song learners” and cites the evidence that young males establishing a territory for the first time appear to be able to crystallize songs that are close matches to those of their neighbor males. How would you respond to this assertion?
I would say that the sparrow most likely socialized with other species in the territory since socialization influences the behavior of the bird (page 49-52). Their songs crystallized when they’re establishing their territory for the first time, but they’re still in the end of the sensory-motor phase and are still overproducing and end up narrowing down their song choices based on their neighbors song.

I believe Hahn described this in OH as they learn songs 1-8 in critical period. Then when they are establishing territory and their neighbors are singing 3 5 and 7 it will crystallize those songs. I would say that the birds needed to have been exposed to those songs during critical period, to be able to crystallize. As far as different dialect?? any ideas beyond each neighbor has a different dialect?  

I think that the dialect evolved based on the environment.  The bird will begin to produce songs based on the acoustics of the environment.  An example is that birds living in the city will produce fast, high pitched songs whereas birds in the country or forest will produce longer, lower frequency songs.

 Define a song dialect, and contrast song dialects with at least two other forms of inter-individual variation in song. Explain what kind of information you would need in order to distinguish among all of these forms of interindividual variation in song.
Song Dialect-Different song types, may be “marked off” in different geological areas. Must occur with distinct markers --> 1. Variation within an area less than variation between areas. 2. Boundaries between song variants abrupt.
The answer key says “A song dialect is a form of inter-individual variation.”
Inter-individual variation (Non-geographic Variation)-song varies only on individual basis, no distinct areas of same song types. Multiple individuals in one area could have multiple different song types.
Clinal Variation (Non-dialectal Geographic variation)- song type gradually changes as you move through an area. Variation in song types in areas next to each other are very difficult to distinguish, but variation in song types of areas at either end of cline more easily distinguished. (further apart, more variation)

 Give one proximate and one ultimate explanation for why the songs of birds living in different habitats (e.g., densely-vegetated versus more open) might emphasize different sound pitches. Do the same for birds living in urban versus wild woodland habitats.
Vegetated vs. open:
-Proximate- Growing up, bird listens to ,say high frequency song and therefore, learn to sing high frequency
-Ultimate-Densely vegetated areas have more barriers to sound, lower frequency sound is transmitted more effectively. Over time, the birds have adapted to using lower frequency for this reason.
Urban vs wild:
-Proximate- Young birds are likely to acquire the song types they can most easily hear, and so urban birds hear high-pitched songs, therefore develop a song in the high-frequency range.
-Ultimate-  Urban environments are filled with sounds in the lower-pitched frequencies.  This is why urban birds use higher-pitched vocals so they can be better heard by potential mates or enemies.
 What are the stages of song learning in male songbirds, and what is going on in each stage?
There are three stages of song learning in male songbirds:
-The first is the sensory phase where the bird must hear model songs to copy during the critical period. The bird will have a strong preference for conspecific song and is very sensitive to the sequence of phrase pairing of models.
-The second stage is the sensory-motor phase where the bird must practice producing songs and be able to hear to match tutor songs. In this phase, the bird over-produces its song (produces a greater range of what they sing).
-The last stage is the crystallization phase where the bird selects song(s) to perfect and sing. Social environment affects song selection from the overproduced repertoire.
 One of the key features of so-called “critical period” songbirds is that memorization of new songs is only supposed to be possible during a brief “critical period” early in life. Imagine that you are studying a species of songbird believed to be a critical period learner, but that you discover that the song repertoire of males tends to increase from one year to the next as the males age. Could this increase in song repertoire with age occur even if the birds really are critical period learners? If not, why not? And if so, how?
Many wild songbirds hear many different songs during their critical period. During their initial crystallization stage, they choose only a few songs to crystalize. However, their brains become more plastic over winter, and then when they get back into the spring season and get new neighbors, if one of their neighbors are singing a song they haven’t crystallized but they had heard during their critical period as a baby, they may decide to bring up that memory and crystalize that song. This means that they have added a song to their repertoire! However, they didn’t learn the song they heard as an adult, the song must have been brought up from memory and crystallized.
 What is a “neural template” for song learning? Describe how you think the “neural template” might function in song learning of an individual male bird that has received tutoring by tape recordings of conspecific males as compared with in an individual male that has received no tutoring at all. A good answer will discuss the hypothesized role of the “neural template” in both motor production and learning.
The neural template is the guiding principle in the nervous system that is responsible for the motor output of the song and how it’s learned. it guides the process of learning and production. In different birds it produces different functions. In the learners it acts as more of a guide during the learning process of songs, but in suboscines like the eastern phoebe it is the entire blueprint of the song because they are non-learners. ....idk about the role of motor production.....
During the sensory-motor phase, the birds refer to this neural template (along with the songs they learned by tutoring if they received it) and correct themselves based off it.
 Which of the following statements would you say better describes the “neural template” for (a) a white-crowned sparrow (an oscine songbird) and (b) an eastern phoebe (a sub-oscine songbird)? 1. The neural template provides a blueprint for what the bird is going to sing. 2. The neural template provides a guide for the learning process. Defend your answer.
(a) A white-crowned sparrow - 2. The neural template provides a guide for the learning process.  → White-crowned sparrows are actual song learners.
(b) An eastern phoebe - 1. The neural template provides a blueprint for what the bird is going to sing. → Eastern phoebes are vocal non-learners.
 Describe some evidence suggesting that female songbirds may use male song characteristics to evaluate the “quality” of the singer.
(Not sure if this is the experiment hes referring to) Tested female attraction to males that were tape tutored. Tested with males that produced good copies of the tape tutor song and males that produced poor copies. Females (wild) were shown to favor males that produced good copies of the tape tutored song.
 Describe the evidence that male white-crowned sparrows do not need to be tutored with complete songs in order to learn to reproduce essentially normal complete songs. What does this tell us about the nature of the “template” for song learning/production and how it influences the nature of songs produced during adulthood?
The slide called “Syntax Assembly from Partial Song Models” chapter 2A. Their neural template is more of a guideline (compared to the phoebe’s full on map). Although only fragment of a song was played during the critical period, the wcs used both its innate and auditory cues they hear from the song to figure out what they should sound like.  
 Describe the evidence that female red-winged blackbirds are more discriminating when it comes to song than males are. Provide one proximate and one ultimate explanation for this fact.
Expt.1 Female red-wing blackbirds tested for attraction displays to song of male red-winged blackbird vs. a mockingbird imitation. Found that females were able to distinguish between the real song and mimicked song-showed more displays to male RWB’s than mockingbirds.
Expt 2. measured male RWB responses to fellow male RWB songs or to mockingbird imitation. Showed that males were less able to distinguish between the two types.
 What is meant by the phrase “interactive theory of development?” Explain how this theory involves both the “internal environment” and the “external environment.”
The interactive theory of development states that phenotype is a product of the interactions between genotype and the environment. “Environment” includes both internal (chemical environment in the body, for example hormones, neurotransmitters, and nutrients) and external (outside the body) components.
 Describe evidence suggesting that changes in worker bee behavioral phenotype are related to changes in gene activation. Explain how this example illustrates the “interactive theory of development.”
The bee’s genotype responds to the environment in ways that influences the development of the bee’s phenotype (which includes proximate mechanisms underlying behavior, such as the nervous system and behavioral traits). Within three weeks, the bee’s phenotype changes from cleaning cells to feeding larvae and nestmates to packing pollen and lastly, to foraging. Studies by Whitfield have shown that comparison of the genes at various phenotypic changes yield changes in genetic activity. This example illustrates the “interactive theory of development” because it shows how the bee’s phenotype is a direct result of its interactions between environment and genotype. If you place a bee in an environment composed of young bees, that bee will turn off the genes for cleaning and feeding faster than normal and start expressing the genes for foraging at an earlier age. Conversely if there are higher than normal proportion of older bees then the bee will clean and feed for a longer period of time (express the genes for younger bee activity for a longer period of time).
 Describe three hypotheses to explain the presence of blackcap warblers (which historically always wintered in sub-Saharan Africa) in Britain during the winter. Which of these hypotheses was favored by the researchers who studied this phenomenon, and how have those researchers managed to narrow in on that explanation? What assumption involving the rules the birds follow for choosing orientation direction in the autumn must be true for the researcher to be able to draw the conclusion they did from the experiment they did? Propose a straightforward experiment that would test for the validity of this assumption, and describe specifically how you would interpret different possible outcomes of this experiment. Hint: You need to deal with the concept of navigation to answer this well.

1. The birds lost migratory abilities (and just stayed in UK year round) → this was ruled out

2. Birds were migrating from Scandinavia- Travel southwest, end migration in UK

3. Birds traveled from Central Europe- would mean that birds were orienting north-west.

 → Expt.) took adult birds wintering in britain. Found they oriented northwest. When breeded, found that offspring also oriented northwest.

 --if birds were from scandinavia, they should have oriented south-west. So this supports hypothesis 3, that the birds came from central europe.

 --also defends assumption that the birds don’t “navigate”. Orient toward a direction, not destination.


 Given that migratory behavior of blackcap warblers has a genetic basis, how would a new population wintering in a new area (e.g., Britain) originate? A good answer to this question will describe a logical evolutionary scenario involving the adaptive significance of migratory phenotypic traits and how it affects evolutionary change in the population.


Genetic variation exists in every population, and has effect on natural selection. (Other genetic effects on natural selection include genetic drift, bottlenecking, etc, but these are probably not playing a role here.) Basically, there are always a few birds who don’t quite follow along with the average thing everyone else is doing. Usually these birds (who migrate to different places) just die and don’t pass on their genes. But every once in a while, they migrate to a place where they can survive and pass on their genes. Consider this a type of directional selection, favoring birds that would potentially respond well to climate change with their flexible genotypic plasticity.
 A graph of “mid parent residual onset of migratory activity” versus “mid offspring residual onset of migratory activity” reveals a positive relationship with substantial scatter around the best-fit line. What is the primary significance of the positive relationship? What do you think is significant about the substantial scatter around the best fit line?

Positive relationship- suggests that genetics play a large role in bird migratory activity. Ex.) Parents with early onset migratory activity tend to have offspring that show early onset migratory activity.

Scatter around best fit line-Suggests that the environment is also a factor. If genetics were the only basis, then there would essentially be no scatter, or at least much less.

Graph also implies that the genotype of the parent does not exactly predict the genotype of the offspring. There is still genetic variation in a population.
 Describe evidence indicating that timing of onset of migratory behavior in blackcap warblers is responsive to selection. How could these experiments have been made even more convincing? Describe another example of an experimental study, using a more tractable study system than the blackcap warblers, that provides much more convincing evidence that genetically heritable behavioral traits of populations can be modified by selection.

Experiment-->used directional selection to select for late onset migratory activity. Graph shows three generations-->each generation shows a later onset.

-therefore: directional selection for later migration onset produces generations of later migrants. Furthering evidence of migratory onset being a genetically heritable behavioral trait.

The mouse study on nest size was more “tractable” because it included a control, many more generations, and multiple trials/ repetitions, which demonstrated the reliability of the results. (The results weren’t due to random chance.)


 Consider the mouse nest-building selection study described in lecture and in the text. Imagine that the results were obtained with all of the mice held on a constant temperature of 20oC. Now imagine that after the 15 generations of selection shown in the figure that a number of individuals from the “high” and “low” selected lineages are tested for nest building behavior at a range of temperatures from 5oC to 35oC. Imagine that the results show that mice from the “low” selected line build bigger nests at 5oC than they do at 35oC, the mice of the “high” selected line build the same size nests irrespective of what temperature they’re held on, and the two lineages build identical sized nests at 5oC. Discuss the implications of these hypothetical results and specifically address the question of whether these results are consistent with the hypothesis that nest building behavior has a genetic basis. Draw and label a figure illustrating the described hypothetical relationship between nest size, environmental temperature and genetic line.
Contradicts the evidence of the nest building having a genetic basis. Suggests that, in some cases, environment also plays a role...
He drew this out during office hours it was like the lower nest one was variable based on temperature - the colder it got the higher they went, but the high nest mice built the same nest regardless of temperature. So OP is right this is showing that environment plays a role. Environment plays a role in both the low nests (via temp) and the high nests (via nutrition for example)
The same mice species and original population, randomly split up and submitted to different directional selection treatments, where after selection they behave differently to same temperature stimulus. This implies there must have been allelic variation in multiple genes making up the nest-building behavior, in the original population. The “high” lineage mice always built the same size nest, which implies the behavior has a stronger genetic influence, like the mice are fulfilling a pre-set genetic program, and have an inflexible genotype. The “low” lineage mice are building larger nests at smaller temperatures and vice-versa, which implies the behavior has a strong environmental influence. Even within the (probably many) genes coding for this behavior, some allelic gene variants are responsive to change, and some allelic gene variants are completely inflexible to change (in environmental conditions).... I think this is a basic theory of behavior but I can’t remember what it’s called........

After the directional selection, the “high” and “low” populations could be considered different genotypes. In that case, these results imply that each population experiences a different “reaction norm,” where some genotypes are very resistant and other genotypes produce a large range of behaviors depending on environmental input.
 Draw a graph illustrating what you think would happen to the mean value and the variability in nest-building behavior in mice following a number of generations of directional selection on the trait. Do the same if the selection was stabilizing.

Variability would increase from the directional selection treatment.

Variability would decrease from stabilizing selection treatment (mean value would even out and be representative of the whole group).
 Which of the following two statements regarding the control of movement behavior of fruit fly larvae is more reasonable, and why? (1) Wandering behavior is controlled by a single gene locus. (2) Inter- individual differences in wandering behavior can be accounted for by differences at a single gene locus. Explain your choice.
Behavior is best explained by an interaction of genes and the environment. However, inter-individual DIFFERENCES can be attributed to just genetics. Therefore, option 2 would be the best choice. Variances in such behavior also attests to this.
 How do coast and inland garter snakes differ in their interest in, and willingness to eat, banana slugs? Consider the hypothesis that experience with slug-derived chemicals in utero accounts for the behavioral differences observed between coast and inland garter snakes. Propose an alternative hypothesis, and make one prediction for each hypothesis that would help you decide which hypothesis is the best explanation.
Coast garter-Love and prefer to eat banana slugs
Inland garter-large aversion to/avoid eating banana slugs
Alternative (must be proximate-developmental)--baby coast snakes learn to eat slugs from their mothers (after birth, not in utero).
Prediction for original-baby coast snakes would already have a preference for slugs, right after birth.
OR, make sure pregnant mothers do not eat any banana slugs. If hypothesis true, these babies would not show a preference for slugs after birth
Prediction for alternative-baby coast snakes would not show a preference for slugs if taken away from mothers right after birth. Would only show preference after “shown” by mothers to eat banana slugs.
 Contrast organizational and activational effects of hormones and give one example of each.
Organizational effects: contribute to establishment of neuro-muscular and morphological aspects, important for behaviors to be expressed
- Usually occur during a critical period (ie, a period when if hormone not present, may not be responsive to it in future)
- Defined as irreversible (like critical period song learning in birds-- always stuck with the effects)
- Tend to be big morphological effects (degree of early exposure to hormones determine whether a structure is present at all, or functional)

Activational effects can happen any time, not just as adults; opposite of organizational
- Immediate causal, short term effects
- Effects not permanent, reversible, no critical period
- purpose is to modulate activity of existing machinery and how it functions (example of testosterone effecting plasticity of bird song)
 Describe how the embryos in a pregnant rat might influence the development of one another through hormones. Would these effects be organizational or activational? What specifically are 0M, 1M and 2M embryos? What role does alpha fetoprotein play in this system?
Babies “soaking” in maternal estradiol. Estradiol normally masculinizes, which you don’t want because need some females-->babies produce alpha fetoprotein to bind and inactivate estradiol
- But does not bind testosterone--> in brain or other target tissues testosterone is locally converted into estradiol and does its business
- Male testosterone goes into the blood surrounding the embryo, and can go into nearby female and affect their brain too
 -0M=2 Female surrounded by 2 other females (0 males)
 -1M=Female surrounded by 1 male and 1 female
 -2M=Female surrounded by 2 males
   -->2M female more masculinized because brains exposed to more testosterone than 1M or 0M females (actually the estrogen thats having the effect, testosterone is the precursor to estrogen)
 What is lordosis? How have studies using lordosis and aggression as behavioral assays been used to evaluate organizational effects of hormones?
Lordosis - when fertile, female rats arch back and deflects tail to side to facilitate copulation with male.
 Cite experimental data to evaluate the following statement: Laboratory rats can be conditioned to perform any naturally-occurring behavior to avoid an unpleasant stimulus, such as a foot shock.
From the notes: “Rats can be readily trained to run or turn in order to avoid a mild electrical shock. Both running and turning are appropriate natural behaviors for rats to display to avoid things like predators or other frightening stimuli. IN CONTRAST, they appear unable to learn to rear up on their hind legs- a behavior naturally used during exploration for food, not for escape from frightening stimuli. This illustrates that certain types of association are much more easily learned than others.”
---> So, this would be false. They can’t be conditioned to perform ANY naturally occurring behavior. Only those behaviors that would normally be associated with an unpleasant stimulus (running or turning).
 . One of your friends says, “I think the difference between Clark’s nutcrackers and other related birds like pinyon jays and scrub jays in ability to learn tasks requiring spatial memory just reflects the fact that Clark’s nutcrackers are generally better learners than those other species, not that there’s anything special about their ability to perform spatial memory tasks specifically.” How would you respond to this?
Studies were done on Nutcrackers, pinyon jays, and scrub jays to test both their spatial memory abilities and non-spatial memory abilities. For the spatial memory task, the nutcracker significantly outperformed the other birds, suggesting a superior spatial memory. On the non-spatial memory task, nutcrackers did not exceed the abilities of the other birds. Comparing the two tests together suggests that the nutcrackers have a specialized spatial memory. If they were “simply better at learning” then they would have outperformed the other birds in both the spatial, and non-spatial memory tasks...but they didn’t.
 In response to your answer to the above question, another one of your friends says, “Well, I think the differences between Clark’s nutcrackers and other jays are really primarily environmentally determined. Nutcrackers are just motivated to do a lot more food caching when they’re young than the other jays are, so they get more experience caching and retrieving caches, so they get better at it.” How would you respond to this? Propose an experiment that would allow you to determine whether or not this concern is justified.
You could capture some baby nutcrackers, before they have had the ability to “experience caching and retrieving”. Test their spatial abilities in the lab. If their spatial memory still seems to be significantly superior to other birds, than this could be attributed to more genetic factors than environmental
 What is the “nutritional stress hypothesis” with respect to song learning in songbirds? Explain how this hypothesis has both proximate and ultimate components.
One thing that could constrain song characteristics to be honest indicators of male quality relates to the fact that specific brain areas are critical to the learning and production of song, and owing to the extremely rapid development of these brain areas (young songbirds have exceptionally rapid growth and development) there is terrific potential for their functionality to be compromised if young males are exposed to any kind of  nutritional deficit at critical development stages. Thus, only males that were well cared for during the important period of very rapid growth and development and/or males that are particularly resistant to disruptive effects of nutritional deficits during that time – i.e., males likely to be “high quality” individuals – would be able to develop brain regions maximally capable of learning and producing songs. In particular, features of song such as how well tutor songs can be copied, or the size of the repertoire of songs that can be remembered, could be negatively affected by exposure to nutritional stress during development.”
Ultimate component- Females will only choose males with a “higher quality” song. Stress during development could affect the quality of song produced by males and negatively affect their reproductive success with females.
Proximate component- Nutrition during development immediately correlates with the quality of song produced by males later on.
 What is the relationship between feather growth and first year song repertoire in great reed warblers, how was this relationship demonstrated, and how does this finding either support or refute the nutritional stress hypothesis regarding song learning in birds?
The longer the feather length when younger, shows that the bird is either resistant to poor nutrition or getting a lot of nutrition. When the birds are all grown up those with the longest feather length also have the most popular song, this supports the nutritional stress hypothesis because the birds with shorter feathers also had poorer songs.
 Propose an experimental study that could test the validity of the nutritional stress hypothesis (Hint:Look at Figure 2.29 on page 56 for an idea.)
Count the number of female precopulatory displays compared for a group well fed/highly resistant to nutritional stress birds versus. birds that were not given good nutrition during development??
 Beyond providing a potential explanation for variation in the abilities of individual male songbirds to copy and produce tutor songs, what is the ultimate importance of the nutritional stress hypothesis? (Hint: It has to do with the question of why female birds rely at least partly on song for choosing mates.)
because it is an honest indicator of the male bird’s overall health
maybe-ULTIMATE importance--> affects reproductive success with females. Acts as an indicator to females of the health of males. Want to choose healthy males for better fitness of offspring.
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