Animal behaviour Flashcards

1
Q

Define behaviour

A

Internally coordinated responses of whole living organisms to internal and/or external stimuli, excluding responses more easily understood as developmental changes

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2
Q

What is an ethogram?

A

Commprehensive list/description of all bahviours of an organism
- requires extensive study

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3
Q

What types of behaviours can be recorded and how are these measured?

A

Events - short duration, frequency event e.g. aggression, vocalisation, yawning, sneezing, mating attempt
States - occur for extended duration, foraging, movement, freeze response, singing, mating display

Can measure latency, frequency or duration to quantify these behaviours.

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4
Q

Pitfalls when studying animal behaviour

A

To draw conclusions, alternative hypotheses need to be tested - can’t just jump to unsupported conclusion e.g. dawn birdsong

Anthropomorphisms - applying human qualities to non-human animals/things e.g. planning in scrub-jay
- Cannot associate human expressions w/ primates

Human perception vastly different to animals - sensory systems. e.g. sexual dimorphism of Pieris rapae under UV light

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5
Q

Describe Tinbergen’s framework

A

Looked at reason for animal behaviour.

Proximate - recent developments in animals beahviour
- mechanism (causation - physiology, morphology, brain)
- ontogeny (individual history/learning)

Ultimate - evolutionary reasons for improved fitness
- adaptive function -> selection
- phylogeny, evol history

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6
Q

Describe Tinbergens study on beewolf wasps

P. triangulum

A

Q: How do wasps navigate to nest after foraging trip?
Obs: wasps fly in circles above nest when leaving to forage
Hyp: they use local landmarks to find nest again

-> wasp followed landmark when it was moved instead of returning to nest site

Evidence supports landmark hypothesis.
^ example of proximate question, deals w/ behavioural mechanism

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7
Q

What are the three conditions that must be met for Darwinian evolution to act?

A

1) Genetic variation
2) Heredity
3) Differential reproduction

These conditions are necessary for natural selection and sexual selection to enhance survival and reproductive success.

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8
Q

What does evolution act to enhance?

A

Individual survival and reproductive success

This is achieved through natural selection and sexual selection.

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9
Q

What is Lloyd Morgan’s Canon?

A

A specialized form of Occam’s razor/law of parsimony
- most simple explanation should be adopted

It suggests that animal behavior should not be interpreted through higher psychological processes unless necessary.

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10
Q

What is the continental schism in animal behavior research?

A

The divide between Europe and North America in research approaches

Europe: Ethology born out simple field observations. e.g. Tinbergen, Lorenz, von Frisch

N. America:
Focus on physiology + comparative lab based psychology, Pavlov’s classical conditioning formed basis for psychological approach -> developed further into behaviourism.
e.g. Thorndike, Watson, Skinner
- Influence of external stimuli on behaviour
- Observable + quantifiable stimuli/response

Each region has different focuses and methodologies.

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11
Q

What is ethology?

A

The scientific and objective study of animal behavior under natural conditions
- Focused on instinct & fixed action patterns (hard-wired)

It emerged from simple field observations by researchers like Tinbergen, Lorenz, and von Frisch.

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12
Q

How did the approaches to animal behavior research in Europe and North America clash?

A

Due to differing focuses on field observations versus lab-based methodologies

This led to significant differences in research outcomes and interpretations.

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13
Q

What does the term ‘NATURE’ refer to in the context of behavior?

A

Ethology - evol basis of bahviour

It is innate, instinctive + pre-programmed by genetic constraints

The genetic and biological factors that influence behavior.

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14
Q

What does the term ‘NURTURE’ refer to in the context of behaviour?

A

Behaviourism - not constrained by evol history

Driven by environment - experience + learning

Refers to the influence of external factors on behavior.

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15
Q

Support for nature argument

A

Lorenz - all species have repertoire of innate, stereotyped, species typical behaviours (fixed action patterns)
○ Determined by specific sign stimuli in the env

Lack - highly territorial robins use red as a sign stimulus (used stuffed replicas)

^ethologists observed animals in nature

Used stuffed robins to identify sign stimuli like the color red.

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16
Q

Support for nurture argument

A

Behaviourists - data driven experiments on learning theory (quantitative) under controlled conditions.

Watson - ‘little albert’ conditioning exp in humans - linked white rats to a fear response

It also considered environmental influences, but lacked understanding of genetics.

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17
Q

What is the contemporary understanding of the nature/nurture interaction?

A

Both genetic and environmental factors interact to shape behaviour - forms a spectrum

  • web weaving in spiders is innate/genetically determined
  • problem solving in primates (apes) is learnt

This view recognizes the complexity of behavioral influences.

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18
Q

Fill in the blank: The MaTcH study found that traits are _______ hereditary.

A

49%

This meta-analysis examined 14.5 million twin pairs.

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19
Q

What shift occurred in the focus of behavioral studies from thw 1970s onwards?

A

From proximate to ultimate approaches

This change led to the formation of behavioral ecology.

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20
Q

What is the use of behavioural genetic experiments?

A

Can help determine where on continuum a behaviour sits.

Twin studies (same genes different env) - MaTCH

Adoption (different genes, same env) - split-clutch exp

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21
Q

Darwins definition fo sexual selection

A

the reproductive advantage which certain individuals have over others of the same sex and species

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22
Q

Types of sexual selection

A

Pre-copulatory:
- intra-sexual (same sec competition)
- inter-sexual (mate choice)

Post-copulatory:
- sperm comp
- cryptic female choice (mate choice)

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23
Q

How does pre-copulatory male-male competition work usually?

A
  • Males gain more than females by multiple matings
    -> competition between males and females choosing between them

Sperm also very cheap to produce

Sexual selection interacts with natural selection to produce trait distributions
- Species showing reversal of ‘classic’ situation can be useful focus of study

Females are highly selective

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24
Q

Drosophila intra-sexual selection

A

Male drosophila get increasing fitness w/ increasing numbers of mates, not so with females - Bateman

  • Females have restricted number of eggs, mating is costly -> so they look mate w/ ‘best’ male
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25
What causes development of sexual dimorphism? ## Footnote Can inc weaponry
Intra- (outcompete rivals) & inter-sexual (be more attractive) selection
26
Give examples fo sexual weaponry
Californian elephant seals 4% males responsible for 85% reproductive success, males fight for dominance so dominant males mate far more often. -> reproductive skew BUT can have trade-off -> horn + eye size in beetles, larger weapons sexual beneficial but reduces survival in Soay sheep
27
What adaptations have arisen to promote inter-sexual selection (female choice)?
Colouration in males, courship displays, elaborate architecture e.g. Bower birds make structures, sticklebacks creates nests
28
What are direct benefits of inter-sexual selection for females?
Oral gifts: nutrients to consume e.g. Great Grey Shrike Seminal gifts: protein-rich ejaculate to nourish female e.g. bush crickets
29
How did female choice evolve?
Sexy son hypothesis - females choose males based on producing offspring w/ the best chance of reproducing Zahavi's handicap principle - females choose males because he was able to survive w/ a costly sexual trait (sexual signals are costly, 'honest signal')
30
What is runaway sexual selection?
Male traits originally indicate survival advantage, genes arise that enable females to detect + chose that trait. -> leads to runaway selection where females prefer longer tails despite fitness cost But no precise genetic basis known for female choice
31
Runaway selection in long-tailed widow birds
Males have multiple mates & multiple nests. Andersson (1982) experimentally manipulated tail length to examine if: - Females preferred males w/ longer tails - Tails were costly -> males w/ artificially elongated tails had more active nests Cutting + sticking tail feathers had no effect, low sample size? Then looked to see if tails were costly -> males w/ shorttails in better body condition over time (handicap principle)
32
What is sperm competition? ## Footnote Post-copulatory
Competition between sperm from rival males to fertilise a females eggs - Extended for of male-male competition - Males have numerous strategies for dealing w/ sperm competition
33
How can risk of sperm competition be reduced?
Mate guarding - avoid competition, common in invertebrates Removing rival male sperm during mating e.g. complex male genitalia remove rival sperm in Odonata Increase investment in sperm production
34
Animal behavioural adaptations for sperm competition
Red Jungle Fowl strategically alocate sperm to maximise reproductive success. - higher investment when presneted w/ novel female Lareg testes: norhtern giant mouse lemur testes 5.5% body vol
35
What ic cryptic female choice?
Females use physical or chemical mechanisms to control male's success in inseminating her eggs. - red jungle fowl females eject more sperm after mating w/ subordinate males (lower ranking), dependent on social standing
36
What are the assumptions made in foraging theory?
1) Animal's goal is to maximise amount of food it can collect per unit time 2) Strategy is series of independent decisions
37
What does the foraging model predict?
Whether an item is included in an animals diet should not depend on how common that item is (no lambda in r2 expression, left), only depends on how common better food types are SO poorer items should be included in the diet only if better items are rare.
38
Can foragin theory be applied to env conditions?
Krebs study into Parus major birds (1977): - Worked with large + small mealworms on conveyor belt - Controlled encounter rate + proportion of good/bad food sources When big prey common, took only big prey When big prey rare, took both big + small prey -> predictions qualitatively & quantitatively correct, birds effective at maximising gain per unit time
39
Does variability impact foraging?
Yes - shown in J. phaenotus birds - Given same number of seeds on average at each feeder, should use them equally? - 1 feeder always gave exactly same number, other gave variable number When avg. number seeds high, bird chose consistent feeder. When avg. number seeds low, birds always chose more variable feeder. If there are diminishing returns of total reward vs fitness - higher tolerance for risk
40
Impact of danger on foraging
Juvenile blue-gilled sunfish are prey fror bass. 1. Adult bluegill foraged open water + grew same rate on both sides 2. Juveniles used open water on bass-free side + foraged in vegetation on side w/ bass 3. Juveniles grew 27% slower in side w/ bass Had to balance foraging habits w/ risk of being eaten (danger) ## Footnote Werner et al, 1983
41
Example of predator avoiding dangerous prey
A. trifolium spider can rush in + bite or wrap it in sil (less risky) Lepidoptera + diptera have probiscus/non-dangerous vs othoptera (crickets) have mandibles (more dangerous) Food type modifies predatory behaviour -> safety prioritised
42
How can learning affect foraging?
P. inorta (finch) have independent foraging patterns depending on tree - theory suggests should be 1 optimal approach SO theory does not take into account ability of bird to learn one behaviour very well -> reducing handling time (complicates theory)
43
Define absolute fitness (W)
Expected number of offspring that an individual will produce over the course of its lifetime. - depends on viability & reproductive success
44
How is relative fitness calculated?
absolute fitness (W) / population mean fitness
45
What is predation?
Predation - interaction where fitness of one organism increases, fitness of the other decreases
46
How do mutualism, competition & commensalism affect both organisms involved?
Mutualism: +/+ Competition: -/- Commensalism: +/0
47
Define crypsis
Ability of an organism to conceal itself especially from a predator by having a colour, pattern, and shape that allows it to blend into the surrounding environment. Can help animals avoid predation
48
Crypsis in moths
Change in selection pressure (industrial rev) -> selection for carbonaria moth (increased frequency) Pollution now reduced so typica selected for again (morph switching) ## Footnote Crypsis dtermined by env slection pressures (predation)
49
Can crypsis include behavioural components?
Yes - resting orientation of moth head on tree affects likelihood of being spotted by predator
50
Define masquerade & compare it to crypsis
Organisms attempt to be mistaken for non-prey, exploits cognitive limitations. vs crypsis where organisms exploit predators sensory limitations (camouflage)
51
What animals can exhibit masquerade?
P. cresphontes butterfly Cuttlefish (cephalopods) capable of both
52
Aposematism
Warning colourations to deter predators due to a toxic trait carried by the organism (honest signal) Can be costly - maintenance + no camouflage so rely on predator learning about toxic trait in past Is maintained in a pop, not selected againts by evol pressures. ## Footnote e.g. black + yellow bees + wasps, brighmonarch butterflies
53
True or false: aposematism is density independent
FALSE - it is density dependent e.g. Spotted bird grasshopper - if individual rare in env, camouflage more beneficial as most predators not aware of toxic trait
54
Examples of aposematism
- Black/white colouration of skunk -> foul smelling spray - orange/black wings of monarch buterfly -> ingested toxic milkweed (similar to cinnabar moth caterpillars) - yello/black of wasps & bees -> sting
55
What did aposematism provide a model for?
Mimicry
56
What are the 2 types of mimicry?
Batesian mimicry - undefended organism mimics well defended model e.g. milksnake benefits (predation as cost for coral snake, left) Mullerian mimicry - both organisms are defended, have evolved similar warnings e.g. hymenoptera (mutualism)
57
What are the differences between crypsis, masquerade & mimicry?
Prey avoids notice in crypsis but is misidentified in masquerade/mimicry (sensory vs congitive systems exploited) Masquerade is a resemblance to an inedible or inanimate object vs mimicry can resemble a trait of another organism to gain a selective advantage SO affects evol dynamics
58
Predator deception in matador bug
Males + females have same size flags + wave at same rate. Unaffected by social context so not for sexual selection. In presence of mantis', flags wave vigorously -> mantis attacks legs, falls off allowing matador to escape. (deflection mechanism/sacrifice) Also thought to be an aposematism to birds (warning signal)
59
Parental care
Behaviour that increases fitness of offspring (likely to have originated/maintained for this function) ## Footnote Rare in arthropods
60
What is the significance of provisioning gametes?
Provisioning of energy & nutrients (proteins + lipids) ## Footnote Larger eggs in some species, such as turtles, provide greater nutrient reserves at hatching.
61
How do some insects provide protection to their eggs?
Coat eggs with defensive secretions ## Footnote beetle species.
62
What are the common materials used for nest building?
Mud, plant material, silk, mucus ## Footnote Nest building can vary from simple burying (sea turtles) to elaborate structures (burrows).
63
What is oviposition site selection?
Non-random egg laying Seen in mosquitoes + dusky warblers ## Footnote Birds and some fish choose specific sites to maximize survival and minimize predation.
64
Give an example of a species that selects oviposition sites to benefit parents.
Female grass miner moths ## Footnote They choose sites that may enhance their own survival.
65
What is egg/offspring attendance?
Parents remain with eggs/offspring at a fixed location -> protection from predators, fanning, site maintenance ## Footnote This form of care is common post-natal, e.g., African bullfrogs dig channel to maintain tadpole water source.
66
What is egg/offspring brooding?
Parents carry egg/offspring -> mobile attendance giving protection. Safer than regular attendance ## Footnote e.g. arachnids (giant water bug), marsupials like kangaroos, mouth brooding cichlids (fish develop in males mouth) -> evolved in many variable environments
67
What is food provisioning?
Offspring provided with a food source after hatching ## Footnote Can be direct (regurgitation in seabirds/beetles, passerines give actual food) or indirect (crowned plovers alert offspring to food source)
68
What are some specialised food source provided by parents?
Milk (mammals), trophic/unfertilised eggs (poison dart frogs) & matriphagy (some arachnids + earwigs) ## Footnote matriphagy - offspring consume mother
69
How do some longer-lived vertebrates provide care after nutritional independence?
Bewicks swans assist offspring in competition for food Burying beetle larvae nutritionally independent at 72 hours.
70
What are the terms for different levels of precocity in offspring?
Superprecocial - independnet living soon after birth Precocial - relatively mature/mobile at birth Altricial - very young/immature, dependent on parental care after birth ## Footnote largely dependent on ecology + env pressures
71
What are major costs of parental care?
Increased predation risk - male pipefish more conspicuous to predators Physiological costs - male cotton top tamarin lose 11% body weight in 3 months >260K to raise a child in the UK
72
What are the benefits of parental care and name the different types?
- Improved offspring survival + quality - Costs & benefits differ between species due to ecology, can differ between sexes. Maternal, paternal, bi-parental ## Footnote Each type varies in its ecological context and species.
73
What mating system is often associated with bi-parental care?
Monogamy ## Footnote Involves a pair bond between male and female.
74
In polygyny, how do mating patterns differ between males and females?
Males mate with several females; females mate with a single male ## Footnote This system is commonly linked to maternal care.
75
What is polyandry?
A female mates with multiple males ## Footnote Often associated with paternal care.
76
What percentage of bird species exhibit bi-parental care?
Oviparous - 90% Most species have altricial chicks - Few sex-specific anatomical/physiological specialisations - Both sexes capable of nest building, incubating, feeding etc. ## Footnote Only 8% have maternal care and 2% have paternal care.
77
What has been argued about paternal care in birds?
It may be a primitive form of care ## Footnote Supported by fossil records showing paternal care in primitive birds.
78
Why do males often provide care in relation to egg costs?
Cost of large eggs to females and cost of care ## Footnote Male care may have originated to protect eggs and attract females to territory.
79
What is the general trend in mammalian parental care?
Females always provide care Monotremes - egg incubation in abdominal pouch, no nipples Marsupials - altricial young cared for in a pouch Placental - wide range of female care ## Footnote However, there are exceptions in certain species.
80
In which mammals do males assist with parental care?
Primates, carnivores + rodents Female care primitive BUT bi-parental thought to evovle 9 times from it ## Footnote The majority exhibit uniparental care, with paternal care being more common than maternal.
81
Parental care in fish
Majority have no care - 78% cases uniparental & paternal more common than maternal inc hiding of eggs, nest building + egg/fry protection Male only care evolved 22 times (evolutionarily labile), always lineages w/ external fertilisation ## Footnote This occurs primarily in lineages with external fertilization.
82
What does natural selection favor in terms of parental behavior?
Behaiour that will maximise lifetime reproductive success (LRS) Sex differences in costs + benefits influence probability of care giving. ## Footnote LRS includes success from current and future broods.
83
Why is maternal care considered common?
Parental investment - sex that invests less in gametes & embryos less concerned about what happens to offspring after mating. Paternity certainty - Males less sure than females that offspring is theirs - males no assurance vs female definite 50% genome - fitness gain from desertion must be greater than from staying ## Footnote The sex investing less in gametes is less concerned about post-mating offspring care. pattern devoid in fish!
84
What is parental investment affected by?
- Gamete sizes - Gamete production cost - Reproductive rate - Competition among sex producing small gametes - Males avoid costly parental care to find more mates
85
Hypotheses for paternal care ## Footnote Particulalrly fish
- Desertion opportunity - Sexual selection - Paternity certainty - Territory establishment
86
Variation of desertion opportunity in different species
Higher in males for internally fertilizing species, but not externally fertilising. - mate guarding + know they are only species to fertilise eggs Higher paternity certainty increasing parental investment ## Footnote -> paternal care in fish
87
What is the most common reproductive strategy among fish species regarding gamete release?
Simultaneous gamete release Some Catfish + Gourami males known to release sperm first ## Footnote 36 out of 46 species studied exhibit simultaneous gamete release, often accompanied by monoparental care.
88
Evidence that paternal care may have evolved via sexual selection due to female choice
- evidence that females prefer nests w/ eggs already in them e.g. mediterranean blenny - preference for high quality territory in Lake Victoria Cichlids
89
Territory establishment in fish
Thought to be selected for by females. Males defend it w/ eggs & fry when externally fertilised species lay eggs on his territory -> lower cost as aldready defending territory - Enhanced paternity certainty - Increased fitness due to egg/fry protection ## Footnote When survival rates are low with only one parent caring, this can lead to the evolution of biparental care strategies.
90
What species have evolved a phenotype to challenge paternity certainty from territory establishment?
Blue gill sunfish Have sneaker phenotype - mature earlier + dart into males territory + fertilise eggs.
91
What is forced adoption in birds?
Occurs in 1% of bird species ## Footnote Forced adoption happens when chicks are raised by parents that are not their biological mothers, often linked to brood parasitism.
92
What are the two types of brood parasitism?
Obligate and facultative ## Footnote Obligate parasites are completely dependent on other species for reproduction, while facultative parasites use parasitism as an alternative reproductive tactic.
93
Maternal care and how it arises in birds
Maternal care + polygyny common in seed eaters e.g. savannah weaver birds. - Seasonally abundant food resource , lower cost to uniparental care - Internal fertilization -> male opportunity to desert ○ Male gains more by desertion (LRS depends more on number of matings)
94
How does biparental care arise?
Constraints on offspring survival w/ uniparental care. Can be simulatanoeus or sequential (emperor penguins divide labour in breeding cycle)
95
What can increase parental co-operation?
Lower sexual selection + sex ratios not skewed (equal)
96
Adoption in birds
In Common eider (Sea Duck), 47% chicks not raised by mothers - likely to abandon young if mothers in poor condition Can be adopted: - Gain parental experience - Low costs - High cost to getting it wrong ○ Could kick out your own offspring ○ Ignore begging from own offspring
97
What is brood parasitism?
Forced adoption - occurs in ~1% bird species. ## Footnote Cuckoos & cowbirds
98
What are the 2 types of brood parasitism?
- Obligate - parasite completely dependent - Facultative - parasitism an alternative tactic + supplements parasites own reproduction
99
Communication between animals
Transmission of information from sender to receiver (animal to animal), receiver modifies behaviour in appropriate way. Can be indirect + eavesdropped Can interspecific & degree of co-evolution between sender/receiver Can be honest (ladybird coluring indicates toxicity) or dishonest (mimicry in moth + flies to look like bees/wasps) ## Footnote e.g. hornet moth, hoverfly
100
Use of pheromones
Chemicals released that infuence behaviour of other members of same species (intraspecific) e.g. kangaroos to detect fertility in females
101
What determines how animals communicate?
Environmental conditions + signal characteristics e.g. chemical signals are long distance but transmitted slowly
102
Evol origins of communication
Many communication components have been taken from other contexts - used for other purposes, often through sexual selection. e.g. in cranes, courtship dance derived from flying
103
Cues vs signals
Signals evolve to increase fitness of sender by altering behaviour of receiver Cues did not evolve this way, despite receiver using them this way ## Footnote Size is a cue, not a sgnal, antlers are signals (costly to produce/maintain)
104
What information can signals convey?
Internal env: - species, age, group membership, kinship - location - condition/quality - fighting ability - motivation/future actions (e.g. attack) External env: - presence of predator OR food
105
What is the benefit of dishonesty?
Low quality males gain large benefit from mimicking high quality males -> mating SO females needs to recognise dishonesty
106
What are honest signals? Give some examples
Reliable signals that are truthful. Costly to send. Cost correlates w/ sender quality. e.g. stotting to demonstrate physical strength in deer - dter predators e.g. male red deer antlers are sign of quality - sexual selection, males also grunt + release pheromones in urine - Larger body, lower frequency - Higher rank, grunt more often
107
Colour and the costly signalling hypothesis
Colour increases predation risk + costly to maintain. -> reamianing alive w/ hindrance indicates strong genetics (Zahavis handicap principle) Colour can also be diet related (good foragers) May also have metabolic tradeoff hindering immune system. ## Footnote Low quality males suffer greater cost than high quality
108
Alarm calls
In blackbirds, can be eavesdropped by other species -> brings attention to predator. Vervet monkeys specialised alarm calls for different predators.
109
How are badges of status used?
In Passerine birds - indicate dominance - honest signal but not conspicuous Artificially enhanced badge in House Sparrow -> 3.62 attack increase by other males -> intra-species social cost, ## Footnote Also in paper waps - have clypeus patch (similar increased aggression when clypeus enhanced)
110
Describe features of dishonest signalling
Up to 44% regenerated claws in Yucca & Yulipes crabs sp. are large but low quality, good for signalling but bad for fighting Dishonesty more prevalent in lower mean body mass populations ## Footnote Hard to detect
111
Sexual mimicry
Sneaker fish found in blue-gilled sunfish, cuttlefish & salmon: males mimic females to fertilise eggs in other dominant males territories - Cost of falsely rejecting mates
112
Aggressive mimicry
Predators can lure in prey -> spider tailed horned viper, anglerfish
113
Define dispersal
Movement of an organism away from birth site or population centre to colonise new areas.
114
Lemming population dynamics ## Footnote D. hudonius
Solitary but come together to breed (prolific w/ 10 pups per female) Short maturation time of offspring so would expect exponential pop growth or plateau due to lack of resources BUT instead has fluctuating pop (Greenland arctic tundra)
115
What factors could influence fluctuating lemming pop dynamics?
Migratory behaviours persist despite no selective advantage to individual (not to own death though) Stoats (M. erminea)? BUT gestation too slow + when lemming pop 'turns' ratio lemmings:stoats is highest, lowest predation pressure SO likely predation aggregation: long-tailed skua, stoat, snowy owl + arctic fox - controls lemming dynamics
116
What did Hot theorise about pop dynamics 1985?
No dispersal in temporally constant environments. -> individual that moves will have slightly lower fitness than individual that stays ## Footnote Due to carrying capacity of a population
117
Costs to dispersal
Energetic costs - fewer eggs laid in D. melanogaster after dispersal Investment in dispersal physiology - winged vs wingless cricket (G. assilimilis), treated group invested repossessed energy from wings into ovary production Predation - voles (M. oeconomus) that dispersed died sooner Failure to find new habitat - selection against dispersal in island like systems, continuous vs fragmented feeding patches in butterflies (deaths from dispersal high in fragmented habitats)
118
What is a major problem with Holt's dispersal theory?
Environments rarely temporally constant so carrying capacities likely to change annually.
119
How can temporal variability be studied for dispersal?
Metapopulations - group of populations linked by small amounts of dispersal. Individual populations often small -> can go extinct (stochastic element), - but often repopulated, can characterise age of population
120
Can temporal variability favour dispersal?
Yes it maintains dispersal Dependent on patch/population characteristics e.g. isolation/connectivity
121
What factors can increase dispersal propensity?
- new invasions - anticipation of env - avoiding inbreeding (depression) Sex-dependent dispersalf favoured by anisogamy
122
New invasions select for dispersal
Cane toad intro in Australia - predators (Iguana) not eveolved to deal w/ its toxic defences Toad spread across Australia - toads w/ longer legs moved farther + are in newest patches, started moving in straighter lines -> speed of movement across Australia increased as dispersal ability of toads increased.
123
Anticipiation of env favours dispersal
S. frugiperda (fall armyworm) - corn pest, very high densities of larvae, larger larvae eat smaller ones (cannibalism) Cannibals better flyers, larger wing/body ratio (dispersal favoured in densely pop environments)
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Dispersal to avoid inbreeding
Varied experience of environment between individuals Larger proportions of recessive alleles in inbred populations - more likely to be exposed to negative selection In Eastern meadow vole, individuals in pop w/ siblings more ikely to disperse to avoid inbreeding.
125
Migration in Fall Armyworms
Very sensitive to temperature, use wind to migrate 400-600km a night. - Each female can lay up to 2,000 eggs -> massive pop increase, mass migration
126
Density and gorup living in bats
Mexican free-tailed bats (Tadarida brasiliensis), very large numbers 40 mill in May/June to intercept moth migration + feed. Females have high metabolic demand -> lactation + high body temp Communities started living under man made structures, juveniles from bridge roosts bigger than juveniles from cave roosts -> disadvantage to live in high density cave.
127
Does the ability to acquire social info promote group living?
Rendell -> copying is beneficial BUT this omits competition Reawrd from patch is shared so crowded patches less beneficial.
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Is it better to copy or to choose patches at random?
If patches are even or changing rapidly then there is little benefit in copying. However if patches are stable but very uneven then acquiring info from other individuals (copying) is good. e.g. rats livie in heterogenous env (garbage is food source) so copying useful vs goats in homogenous env so copying not beneficial ## Footnote Smolla et al 2015
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Can group behaviour avoid predation?
NO Groups of prey will attract predators e.g. Brown bear intercepts salmon when aggregated during migration Predation makes group living (density costly) -> red spider mite attacked by predatory mite
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What 4 reasons could explain why prey group together to avoid predation?
Selfish herd theory Predator satiation Reduced vigilance Predator confusion
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What is the selfish herd theory? ## Footnote Enhances group living
Herding together minimises individual risk of predation, not beneficial to the wider group but the individual instead - as group more conspicuous. e.g. Redshanks more commonly targeted by Hawks were furthest from nearest neighours ## Footnote Hamilton 1971
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What is predator satiation? ## Footnote Enhances group living
Magicicada sp. eggs hatch + nymphs burrow into ground around tree roots + remain for ~17 years. Very little interbreeding between broods - spatially distinct. If cicada at very high densities, birds will gorge themselves + rest of high density group will survive - birds can only eat fixed amount. (skins + wings counted) ## Footnote 17 year cycle prevents predator aggregation
133
How does reduced vigilance promote group living?
Can spend more time eating than looking around for predators. Supported by scan frequency + group size in Goa & Blesboks HOWEVER increased foraging time may be due to increased competition for food -> reduced vigilance cost fo living in group
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What is the class that includes songbirds?
Class Aves ## Footnote Includes approximately 4000 to 9000 species.
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What is a call in bird vocalization?
Short and simple vocalization that signals flight or danger, produced throughout the year.
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What is a song in bird vocalization?
Long and complex vocalization produced during breeding season. ## Footnote More than one version of a song type is a repertoire
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What are the main elements of song structure?
Notes/elements: simple continuous narrow frequency band, complex frequency amplitude Syllables: 2 or more notes clustered together Phrases: 2 or more syllable clustered together - can also be series of singe notes
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What does syntax refer to in birdsong?
Specific ordering and timing of different elements.
139
What tool is used to measure birdsong?
Spectrograms.
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What factor influences song variation in the white-crowned sparrow?
Diverse habitats leading to different dialects between areas.
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Which birds typically sing?
Usually males, due to competition to breed. Both female choice + male/male competition have driven song evolution ## Footnote Twin selectuve pressures
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What song characteristics correlate with breeding success in barn swallows?
* Long rattle (more testosterone) * Low frequency (better condition)
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What is a signal of quality in male birdsong?
Complex song. - difficult to fake (quality signal) - may idiciate age/maturity ## Footnote Correlated w/ mating success in great tits, great reed warblers + song sparrows.
144
What information can be contained within birdsong?
* Presence of potential mate * Individual identity * Birth and raising location * Physical location * Territory ownership * Willingness to breed * Condition * Early environmental experience
145
What is subsong in young birds?
Rambling sounds variable in time and pattern.
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What is plastic song?
Assumes posture and produces sounds in discrete clusters with detectable temporal patterning.
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What is crystallized song?
Full song expressed with normal variation in volume, duration, and syllabic structure.
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What are the phases of song learning?
* Sensory phase * Sensorimotor phase * Transition to crystallized song
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Define 'sensory phase' in song learning.
Auditory experience where the song must be exposed to and committed to memory.
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Define 'sensorimotor phase' in song learning.
Involves vocal practice and includes both subsong and plastic song stages.
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What are the types of song learners?
* Seasonal closed learners * Age-limited learners * Open-ended learners
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What did Thorpe discover about birdsong by studying chaffinches? ## Footnote Used a sonogram
Innate predisposition to learn songs of conspecifics, limited learning window using auditory stimuli. Song genetically encoded but environment + learning significantly enhances
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What unique song feature do White crowned sparrows exhibit?
Regional variation in song stages at different ages. Coast vs Montane birds. -> ecology/habitat of bird impacts on birdsong learning
154
Which two species can hear each other's songs during the sensory phase?
Swamp sparrow (M. georgiano) & Song sparrow (M. melodio)
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What bird combines song models of more than 1 species?
Swamp sparrows combine models of both species - learning involves soft/plastic template - When crystallizes, decrease in syllable sounds (stabilisation VS in white-crowned sparrows - adult song does not contain heterospecific components.
156
What role does testosterone play in male birdsong?
Testosterone undergoes seasonal changes. Inhibits plasticity, functions to crystallize songs. ○ If administered early, will cause premature crystallization ○ In castrated males, songs never crystallize
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What is the function of the song production nuclei?
They are involved in the neural circuitry of song production. - Higher vocal centre (HVc) - Robust nucleus of archistriatum (RA) - Tracheosyringeal portion of hypoglossal nucleus
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Seasonal/hormonal influences on birdsong
- testosterone injected females will sing + increased vol of HVc & RA - adult male canaries HVC & PA are 99% & 76% larger in spring than in autumn. - neurogenesis in HVc of adult canary when learning new seasonal song
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True or False: The lateralization of birdsong is linked to specific brain regions.
True When HVc lesioned, cannot produce WT song
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What discrete brain regions are interconnected for learning birdsong?
- Area X - Medial portion of dorsolateral thalamus (DLM) - Lateral portion of magnocellular nucleus of the anterior neostriatum (LMAN)
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What species exhibit high and low sex bias in birdsong
High: sebra finch + canary Low: bay wren + buff-breasted wren ## Footnote Correlates to volume of HVc
162
What is cooperative breeding?
Social system where individuals provide care for offspring that are not their own at cost of their own reproduction. ## Footnote Examples include meerkats (single breeder in large colony), starlings (sibling help raise young), and naked mole rats (queen + subservient colony).
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Proportion of vertebrates that exhibit co-operatve breeding
~300 species of birds (3%), ~120 species of mammals (2%), ~30 species fish (0.1%) - But extensively studied Long-tailed tits are co-operative breeders in UK. ## Footnote Approximately 300 species of birds exhibit cooperative breeding.
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What is a characteristic feature of cooperative breeding systems?
Usually 1 male + 1 female breeders with 1 or more non-reproductive helpers. Helpers can be related or unrelated to the breeders.
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How does kin selection relate to cooperative breeding?
Helping evolved via kin selection where helpers help relatives, leading to inclusive fitness benefits. ## Footnote This ensures that helpers are more likely to assist those they are genetically related to.
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What impact does promiscuity have on cooperative breeding?
Promiscuity reduces offspring relatedness and certainty of offspring. ## Footnote Lower promiscuity leads to higher relatedness and more common helping behavior.
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List some shared characteristics of cooperative breeders.
Life history constraints: * Delayed maturity * Low adult mortality * Non-limited dispersal * Low reproductive rate Ecological constraints: *Territory shortage * Mate shortage * High dispersal cost * Low independent breeding success ## Footnote These factors contribute to the likelihood of cooperative breeding in a species - helpers have littlpe opportunity to breed on their own.
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What is the significance of the habitat saturation model in cooperative breeding?
Seychelles warbler intorduced to new island. Cooperation started only after all territories were taken. ## Footnote This model shows that cooperative breeding can emerge when individuals have no other breeding options.
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What happens with the superb fair wren when there arent enough mates?
- Both breeders removed -> 0% male helpers moved - Only breeder male removed -> 96% male helpers moved - Mix of cooperative groups & monogamous pairs, ## Footnote This indicates the dependency of helpers on the presence of breeding pairs.
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What is the 'pay to stay' theory in cooperative breeding?
Helpers work harder for refuge access, which provides direct fitness benefits. ## Footnote This theory highlights the trade-offs helpers make for group membership.
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What is the indirect fitness benefit for helpers in cooperative breeding?
Benefit by helping relatives - close relatedness increases help provided. Increases propagation of related alleles. Long tailed tit: failed breeders often become helpers, 94% chose to helps nests of relatives
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What is the effect of removing helpers?
Significantly reduced offspring raised and food brought in Florida Scrub Jay and African cichlid fish. Increased food due to helpers and not territory quality. ## Footnote This experiment demonstrates the importance of helpers in reproductive success.
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Fill in the blank: In cooperative breeding, helpers often gain _____ experience which enhances their breeding success.
breeding ## Footnote This experience can lead to improved reproductive success when they eventually breed.
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What factors determine the participation of helpers in reproduction?
* Level of relatedness * Competitive ability * Constraints on independent breeding * Reproductive skew models i.e. contest ## Footnote These factors influence whether helpers can participate in breeding.
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Territory/mate inheritance
Direct benefit of helping. Clownfish - local helpers improve chance of taking over territory if vacancy opens
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Reproductive concessions
Breeders offer incentives to helper to stay, more required if unrelated or lots of good territories around e.g. acorn woodpecker - breeders concede more reproduction when more vacancies available
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Reproductive restraint
Helpers opt out of reproduction, IF only relatives around or serious breeding punishments e.g. meerkats - females breed more when unrelated males in group
178
Why live in groups?
Resource and predation related factors ## Footnote Group living can help with resource availability and reduce predation risks.
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What is the trade-off in group living?
Increased competition vs predation risk ## Footnote A balance is required to maximize fitness in social groups.
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What did Markham et al 2015 find in Amboseli baboons?
Predation risk decreases & competition increases with group size ## Footnote Intermediate group sizes are associated with higher fitness.
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What variables were accounted for when modelling optimal group size in baboons? ## Footnote Markham (2015)
Fecal glucocorticoids (stress hormones), daily travel distance ## Footnote These variables help assess stress and resource availability.
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What leads to conflict within groups and potential social hierarchies?
Resource competition - Dominance hierarchies occur where individuals can monopolise resources - More egalitarian groups occur where individuals cannot monopolise (e.g. Howler monkeys), small groups more successful
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What is the relationship between rank and stress in baboons?
Lower rank = higher heart rate and blood pressure ## Footnote Stress is linked to health and elevated glucocorticoids.
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What happens to baboons after dominant males are removed?
Group becomes less aggressive; HR/BP decreases ## Footnote This indicates that social structure affects individual stress levels.
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What is scramble competition?
Competition where individuals compete for resources without direct confrontation ## Footnote It leads to decreased immune function and reproduction.
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What are coping mechanisms to reduce aggression in groups?
Intervention and reconciliation ## Footnote Social behaviors can help manage stress and conflict.
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Which species experiences the highest stress changes by rank?
Barbary macaques ## Footnote Mid-rank individuals are often the most stressed.
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What is the social brain hypothesis?
Increased neocortex ratio correlates with group size ## Footnote This suggests cognitive demands increase with larger groups.
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What annimals form social bonds (friendships)?
- In Kaimanawa horses in NZ - females more integrated in their social groups have better reproductive success than those that are poorly integrated. - In Amboseli baboons - females that have stronger bonds live longer & have more babied. ## Footnote Females with stronger social bonds have better reproductive outcomes.
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What is the role of matriarchs in Käller Whales? ## Footnote Reproductive cessation
Matriarchs remain together + form strong bond, improve foraging success of the group ## Footnote Their presence enhances group survival and resource acquisition.
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What happens in elephant groups with younger matriarchs? ## Footnote Reproductive cessation
Less well adapted to distinguishing between male and female lions ## Footnote Older matriarchs have more experience, enhancing group safety.
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What is associated with grooming in social relationships?
Release of endorphins ## Footnote Reward centres pirmed by social behaviour.
193
What is the annual distance traveled by sooty shearwaters during migration?
64,000 km ## Footnote This distance highlights the extensive migratory patterns of certain bird species.
194
Define migration in the context of animal behaviour.
Regular movement back and forth between two relatively distinct locations by animals that use resources concentrated in these different sites ## Footnote Reference: Alcock, 2013.
195
What are the three types of orientation important for migration?
* Pilotage - steering a course using a familiar landmarks * Compass orientation - ability to head in a particular compass direction without reference or landmarks * True navigation - ability to orientate toward a goal such as home/breeding area/ without use of landmarks + regardless of its direction e.g. Reed warblers ## Footnote These orientations help animals navigate during their migratory journeys.
196
How many times has migration evolved in Catharus thrushes?
Three separate times or alternatively has been lost from all other species ## Footnote This indicates the complexity of evolutionary adaptations related to migration.
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List some costs associated with migration.
* Extra weight must be carried to build sufficient energy reserves * Temporary atrophy of reproductive organs * Increased muscle contraction efficiency * Altered metabolism enabling fat storage * Risk of death during the trip * Need to be well adapted to multiple habitats ## Footnote These costs can significantly impact the survival and reproductive success of migratory species.
198
How do energy reserves influence migration routes in red-eyed vireos?
Those with ample energy reserves took a direct route over the Gulf of Mexico, while those with low energy took an indirect route ## Footnote This illustrates the importance of physical condition in migratory decisions.
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What are the benefits of migration ?
- In N. America summer, abundance of protein-rich insects + longer day lengths than tropics - enhanced feeding for birds - Cyprinid fishes migrate over winter to low predation habitats when feeding opportunities are reduced - Monarch butterflies migrate to Mexico to roost over winter - too cold in N. America so will freeze ## Footnote Migration increases fitness
200
What does Emlen's funnel measure?
Used to measure pre-migratory restlessness in birds and their orientation in a particular compass direction depending on the time of year ## Footnote This experimental setup is used to study migratory behavior in the lab.
201
Name some cues for animal navigation.
* Visual * Olfactory * Atmospheric pressure * Sound * Geomagnetism ## Footnote These cues provide critical information for animals during migration.
202
How is the sun used as a compass by animals?
At a given time of day and year, the sun's position provides compass information ## Footnote This method is utilized by various species, including starlings.
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What did Kramer (1951) demonstrate regarding starlings?
Starlings use the sun for navigation and those with phase-shifted endogenous diurnal clocks misinterpret the direction of the sun ## Footnote This study highlights the importance of circadian rhythms in navigation.
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Which animals have been shown to use the sun for navigation?
* Frogs * Butterflies * Turtles * Lizards * Rodents ## Footnote This suggests a widespread adaptation for using solar cues in navigation across species.
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What evidence suggests that pigeons are sensitive to pressure changes?
Pigeons showed a higher percentage of getting lost when olfaction was impaired compared to the control group ## Footnote This indicates that olfactory cues play a significant role in their navigation.
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What navigation mechanism do green turtles use?
They can find their way back to Ascension Island if released north but not south, although the mechanisms are unclear ## Footnote This suggests complex navigation abilities that are not yet fully understood.
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What early experiments found no evidence in pigeons?
Experiments found no evidence of sensing magnetic field in pigeons ## Footnote This suggests that initial hypotheses about pigeons' magnetic sensing capabilities were incorrect.
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What evidence supports that birds have a magnetic sense?
Data from Emlen's funnel gives evidence that birds have magnetic sense ## Footnote Emlen's funnel experiments demonstrated the ability of birds to orient using geomagnetic cues.
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What is a significant feature of Emlen's funnel?
It includes an ink pad, wire screen, and spring ## Footnote These components are part of the experimental setup used to assess bird orientation.
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Where was the magnetoreceptor located in pigeons discovered in 2004?
In the ophthalmic branch of the trigeminal nerve in the pigeon bill ## Footnote This discovery highlighted a physiological basis for magnetic sensing in birds.
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What happens when the right eye of captive European robins is covered?
They cannot orient using geomagnetic cues ## Footnote This suggests that the magnetic sense is localized to the right eye.
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What do migrating insects use to navigate in the autumn?
Geomagnetic cues ## Footnote This indicates that not only birds but also insects have evolved mechanisms to navigate using Earth’s magnetic field.
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What compass do monarch butterflies possess for migration?
Inclination magnetic compass ## Footnote This adaptation aids their autumn migration over long distances.
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How do songbirds exhibit migratory restlessness?
They orient to stellar cues at night ## Footnote This behavior is critical for their navigation during migration.
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What happens to songbirds raised without night sky experience?
They could not orient properly for migration ## Footnote This implies that exposure to the night sky is crucial for developing migratory skills.
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What did Swainson and Gray-cheeked thrushes do when local magnetic cues were manipulated?
They calibrated using cues at twilight ## Footnote This shows that birds can adjust their navigation based on available environmental cues.
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What is path integration used by desert ants?
A navigation method to calculate their position relative to a starting point ## Footnote This method allows ants to return to their nest after foraging.
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What do bats use to calibrate their internal magnetic compass?
Polarised light ## Footnote This ability is also seen in various species including birds, fish, reptiles, and arthropods.
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How do dung beetles navigate?
They orient using polarised moonlight ## Footnote This adaptation helps them find their way in the dark.
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What is required for landmark navigation in animals?
Individuals must constantly record and report information ## Footnote This is essential for effective navigation in complex environments.
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How did adults and juveniles differ in migratory behavior when moved across the USA?
Adults compensated for the displacement; juveniles flew blindly south ## Footnote This suggests differences in learned migratory behavior between age groups.
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What does the migratory behavior in white-crowned sparrows suggest?
That the continental map is learned in these birds ## Footnote This reflects the importance of learning in developing navigation skills.
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What is cooperation?
Cooperative behaviours involve an individual paying a cost to give a benefit to a recipient, increasing their fitness.
235
What should natural selection favour according to the text?
Natural selection should favour genes that increase individual fitness, leading to selfish behaviours dominating over altruistic ones.
236
What is the 'tragedy of the commons'?
It refers to a situation where everyone would benefit from increased cooperation, but individuals gain massive benefits from being selfish, leading to widespread selfishness.
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What are direct benefits in the context of cooperation?
Direct benefits occur when the reproductive success of the cooperator is increased.
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What is direct reciprocity?
Direct reciprocity requires repeated interactions where both individuals involved have increased fitness and memory of past interactions.
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Give an example of direct reciprocity.
Cleaner wrasse cleans parasites from larger fish; wrasse gets a meal while larger fish have ectoparasites removed.
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What are indirect benefits in cooperation?
Indirect benefits occur when the reproductive success of individuals sharing the cooperators' genes is increased.
241
Define altruistic behaviours.
Altruistic behaviours involve a cooperator paying a cost with no self-benefit to fitness.
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243
Give an example of altruistic behaviour.
Vampire bats share blood with individuals not successful in finding food, typically between pairs or relatives.
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What is the significance of repeated interactions in cooperation?
Individuals cooperate on the first interaction, and in subsequent interactions, they copy the behaviour of the other individual.
245
What is Hamilton's rule?
Cooperative genes should increase in frequency when benefits (B) of cooperating are greater than costs (C), considering the level of relatedness (r).
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What does the equation rB - C > 0 signify?
It signifies that cooperative behaviour is favoured when the benefits of cooperation outweigh the costs, adjusted for genetic relatedness.
247
What is kin selection?
Kin selection is the process by which traits are favoured due to their beneficial effects on the fitness of relatives.
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Fill in the blank: Animals must be able to ______ to benefit from kin selection.
recognise relatives
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What are reputational benefits in cooperation?
Cooperators gain reputational benefits, making altruism worthwhile in the long term.
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True or False: Altruistic behaviour can only be explained evolutionarily through immediate self-benefit.
False
251
What role does memory play in direct reciprocity?
Memory of past interactions is crucial for recognizing and reciprocating cooperative behaviour.
252
What dietary strategies can tiger salamanders develop?
Tiger salamanders can develop as cannibals or invertebrate feeders.
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254
How were tiger salamander larvae grouped in experiments?
Larvae were divided into groups with: 1) Siblings only, 2) Siblings and cousins, 3) Siblings and non-kin, 4) Siblings, cousins, and non-kin.
255
What is the mean age of cannibal morphology expression in tiger salamanders?
The mean age (days) of cannibal morphology expression varies with different levels of relatedness.
256
How does relatedness influence shoaling behavior in guppies?
Full siblings spent more time shoaling together and swam closer together than half siblings.
257
What are possible mechanisms for kin recognition?
1) Environmental cues: familiarity or shared environment, 2) Phenotype & self-referent matching, 3) 'Green-beard' effects.
258
What is an example of kin recognition in long-tailed tits?
If nest mates are switched, non-related birds treat each other as relatives, learning who should be relatives, not who is actually related.
259
What did the study on golden hamsters reveal about kin recognition?
Non-kin were investigated sooner and longer, indicating a self-referent mechanism.
260
What is an example of cooperation leading to inclusive fitness?
Courtship in turkeys where males court alone or in coalitions with a dominant and subordinate male.
261
What is the net benefit of subordinate males in turkey courtship?
The net benefit of a subordinate male is +1.7 offspring.
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What is Hamilton's rule in relation to dominant males?
When put into Hamilton's rule: (B-C) = (0.42 - 0.9) = +1.7.
263
What behaviors can infer intelligence in animals?
Planning and use of tools ## Footnote These behaviors indicate problem-solving and innovation abilities.
264
Which animals are primarily studied for intelligence?
Corvidae family (crows, jays), parrots, cephalopods, great apes (chimps) ## Footnote These groups have shown significant cognitive abilities.
265
What is one method of measuring intelligence in animals?
Using brain size as a proxy ## Footnote Larger brains in carnivores are often associated with better puzzle-solving skills.
266
What is an example of social learning in animals?
Blue tits pecking through foil caps of milk bottles ## Footnote This behavior is learned socially rather than genetically encoded.
267
What does the social brain hypothesis suggest?
Social complexity drives the evolution of intelligence ## Footnote It posits that challenges of social living lead to increased cognitive abilities.
268
How does group size relate to brain size according to the social brain hypothesis?
Group size correlates with relative neocortex size ## Footnote This correlation suggests that larger social groups may require greater cognitive skills.
269
What factor predicts brain size when controlling for phylogeny and body size?
Diet rather than group size ## Footnote This complicates direct comparisons of intelligence across different species.
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What does self-recognition in animals indicate?
Awareness of self or consciousness ## Footnote This is often tested through mirror recognition tests.
271
What significant finding did Gallup's 1970 study reveal about chimpanzees?
Higher levels of mark-directed responses with a mirror ## Footnote This suggests that self-recognition evolved in apes but not all primates.
272
Which other species besides apes have shown signs of self-recognition?
Dolphins and Asian elephants ## Footnote These species also exhibit behaviors indicative of self-awareness.
273
What behavior did magpies exhibit in front of a mirror?
Strong responses to gold badges but not to black ones ## Footnote This indicates a level of recognition and distinction in their perception.
274
What behavior was observed in cleaner fish (L. dimidiatus) after mirror observation?
Throat scraping behaviors after a marker was added ## Footnote This behavior suggests recognition of self in a way that resembles parasite interactions.
275
What notable change in behavior was observed in manta rays in front of a mirror?
Behaviors changed, but no mark was applied ## Footnote This suggests some level of self-awareness, though evidence is less direct.
276
Theory of mind
Thought to be uniquely huan - awareness of different perspectives/feelings. Scrub jays try to avoid other birds seeing them store their food - apparent intelligence or theory of mind? Testing in animals 2 containers - grasped or accidentally touched -> primates interested in intentional behaviour. BUT may just be good observers of hand gestures - Primates can use info about environmental constraints
277
Fairness/equality in animals
Economists argue that we are self-interested (game-theory models) BUT data support humans have a moral framework + show inequality aversion. Capuchins show awareness of inequality - observes other individual being fed grapes not cucumber after same task.
278
What is a mating system
Reproductive behaviour among sexually active males & females -> evolutionary differences between monogamous + polygamous animals
279
What 4 factors affect mating systems?
Differential investment - sex that invests more is limiting resource -> polygyny favoured - polyandry occurs in poor quality env (co-op breeding) Ecological factors - if resources are clumped + defensible -> high polygamy potential Paternity certainty - can lead to monogamy if males guard mates Dependence of young - parental care required vs reproductive success for increasing fitness dtermines whether it is better to have more mates or not
280
What can arise due to mating systems?
Secondary sexual characteristics -> intra & inter-sexual competition. e.g. bright plumage of male birds, the antlers of male deer, or the manes of male lions
281
Polygyny
A pattern of mating in which a male animal has more than one female mate
282
Evolution of polygyny
Female defence, resource defence, scramble competition, lek.
283
Resource vs demale defence in zebras
Grevy's zebra defends areas where females congregate Pains zebra defends females from rival males - females congregate (anti-predation)
284
Scramble competition
Females & resources not always defendable. If females widely dispersed, males scramble to find as many mates as possible. e.g. male thirteen-lined ground squirrel search for mates over few weeks of mating period
285
Lekking
Associated w/ extreme traits: e.g. Hammerhead bat evolved large cavern in head, males congregate around trees + make loud noise -> attracts females Lek - aggregation of males that display to females, females do not gen any resources (only there to mates)
286
Benefits of lekking
Males increase chances by having extreme sexually selected phenotypes/behaviours. Females have a pick of males -> can source high quality genes. Unclear for subordinate males - one dominant male white-bearded manakin had 75% of copulations ## Footnote Reproductive success skewed in leks. Subordinate males thought to be younger
287
Why do males bother to form leks?
Hotspot hypothesis: males cluster around routes or places frequented by females. Ev: 4 bird species (2 manakins, flycatcher + hummingbird) in Costa Rica used same location to lek -> topology determines female location Hotshot hypothesis: subordinate males gather around attractive dominant males. Ev: Removing dominant male European Sandpiper -> subordinates dispersed, but subordinate removal did nothing. Female preference hypothesis: males cluster because females prefer sites w/ larger groups of males Closely linked to other 2.
288
Which animal groups demonstrate monogamy?
3% mammals, 90% birds, ~1% fish
289
Mate guarding hypothesis for monogamy
Monogamy is adaptive (more offspring) when females left by one male readily mates w/ another - Beneficial when females remain receptive after mating + difficult to locate e.g. crown shrimp - Better to keep what you have than go looking for another inaccessible female
290
Mate assistance hypothesis for monogamy
Monogamy adaptive when parental care greatly increases offspring survival, often in altricial species. e.g. reproductive success of californian mouse higher if he stays to help rear young
291
Female enforced monogamy
Burying beetles, male + females bury carcass which females lay eggs on. Males try attract another female using pheromones, but 1st female detects these + attacks male -> he stops releasing pheromones. - Nature of resource grants female control over mating
292
Social monogamy in birds
- Egg in nest balances relative investment of both males & females - Rearing altricial offspring requires extra parental care ## Footnote But most species are promiscuous (genetical tools) - engage in extra-pair copulations (EPCs) despite social monogamy
293
Define learning
Changes in behaviour following experience that are not due to maturational growth processes, fatigue or sensory adaptation -> mechanistically is strengthening of a synapse
294
Habituation
Animals stop responding to repeated stimuli - no reinforcement Kandel used repeated stimulation in aplysia (sea slug) via tactile stimuli. -> less effective Ca2+ channels ## Footnote Non-associative
295
Sensitisation
Simultaneous presentation of touch & shock, animal becomes highly sensitive to touch - same network for that of habituation. e.g. Rats behaviour shifts after delivering offspring (parturition) -> maternal behaviour, more likely to pick up pups - sensitisation response to pup odour
296
What is an engram?
Physical basis of memory, due to patterns of synapses e.g. olfactory memories in insect brain stored in muschroom bodies
297
What did Bence study?
Genetic basis of circadian rhythms, learning & phototaxis Dunce mutants in flies had learning + memory deficiencies - odour/shock conditioning experiment Dunce controls recycling of cAMP -> AMP - important for memory/learning pathways e.g. ARM ## Footnote ARM - anaethesia resistance memory Gave behavioural, anatomicla + biochemical insights + fits w/ aplysia data
298
Pavlov's classical conditioning experiment
Salivation responses to bell ringing/feeding in dogs -> associative mechanism
299
Classical conditioning in Blue Gourami fish
Males defend territory from other males. Experimentally conditioned males by announcing arrival of gravid females (about to lay eggs) using buzzer. Conditioned fish -> less aggressive, built nests more, spawned quicker (reduced competition) & produced more offspring - fitness advantage to learning
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How can learning be considered a force of evolutionary change?
Aposematism e.g. harlequin ladybirds, monarch butterfly caterpillar produces alkali (toxic) - Predator learning (associate colour w/ foul taste/toxicity) has led to selective advantage of aposematism - This preference not found inherently in birds -> learning indirectly led to mimicry
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Behavioural adaptations from learning
Bombardier beetle relies on defence (chemical release) upon predation by frog - relies colouration & learning ability of its predators to avoid predation
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One trial learning
Animals link stimulus to response after 1 trial. Taste & nausea linked easily, sound & shock also linked easily - but little crossover between 2 pairs - Selective benefit of specific associations (avoiding toxic foods OR predation)
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Operant conditioning
In lab - Skinner box gives active form of classical conditioning w/ food vs electric shock In field - blue tits pecked holes into foil caps to access cream -> spread through pop via social learning/copying
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Examples of social learning
Honeybees - trained to complete tasks for food reward - w/ human demonstrator ~99% bees learnt to complete task Macaques in Japan wash sweet potatoes left on beach by researchers - spread through pop by social learning, younger animals learn better than older
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Culture in social animals ## Footnote Social learning (especially when young)
Non-genetic inheritance of behaviours, customs + norms. No purpose/benefit for Chimp to put grass in ear but copied by others (persisted)
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What species is insight learning mainly seen in?
Sultan chimp tied 2 sticks together to retrieve food. Crates stacked to reach bananas Vs pigeons which can show beahviour but no insight used (needs to be pre-learnt)
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Spatial learning in Jays
Store seeds, successful in spatial taks but no non-spatial SO learning context specific to animals ecology -> may indicate some fitness cost to learning
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Cost of learning in drosophila
More you learn, quicker you die -> strongly suggests cost of learning in drosophila
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# [](http://) Behavioural definition of sleep
Rapidly reversible state of immobility & greatly reduced responsiveness, homeostatically regulated such that lost sleep is made up with an increased sleep drive. ## Footnote (Siegel, 2008)
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Electrophysiological definition of sleep
Pattern of specific brain wave activity, associated w/ overall decrease in neuronal activity -> NOT global turning down - Series of precisely controlled brain states - Sequence determined by activity of specific brain nuclei
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Why do we sleep?
Basic homeostatic need that increases w/ time awake. - duration + pattern changes w/ age - bout duration increases w/ organism size - critic for normal cog function (rodent fatality w/ no sleep after 2 weeks)
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What factors influence a species sleep patterns?
Ecological: - predation - mating - thermoregulation - need to forage Intrinsic: - memory consolidation - energy conservation - metabolic clearance
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How do we measure sleep?
Electroencephalogram (EEG) - provides continuous recording of brain activity, important tool in lab and clinic -> localises epileptiform activity, mapping - Cheap - Non-invasive ## Footnote Categorise cognitive sleep states using amplitude & period/frequency
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What are the different EEG sleep states?
- Alpha: most prominent when eyes closed - Beta: associated w/ being alert, attentive & actively thinking - Stages 2-4 are slow wave sleep (SWS) or non-rapid eye movement (NREM) - REM sleep is also known as paradoxical sleep -> high freq, low amplitude, associated w/ dreaming
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Describe sleep stage progression
EEG recordings showed sleep consists of number of stages that occur in characteristic sequence. 1st hour is NREM followed by REM: - SWS/NREM & delata activity (>50% deep sleep) - REM ## Footnote Kleitman & Aserinsky
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How many sleep cycles per night?
5 - REM duration increases/SWS decreases throughout sleep bout - deep sleep (IV) only in 1st 2 cycles ## Footnote SWS - slow wave sleep REM - rapid eye movement
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What other ways can we detect the stage of sleep cycle?
Eye movement - electrooculogram (EOG), mainly during REM Neck movement - electromyogram (EMG), waking + REM transitions Heart rate & respiration peak during REM
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Rat vs Human EEG
Rats nocturnal so sleep in day, shorter boyts of sleep + more frequent transitions wake -> NREM -> REM
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C & S model
Interaction between homeostatic sleep pressure & internal circadian clock Circadian component suppresses sleep drive in afternoon to maintain wakefulness.
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What neural systems are involved in sleep?
- Forebrain system independently supports SWS - Brainstem systems activate forebrain into waking & trigger REM sleep ## Footnote Identified by nerve transection studies (Bremer) in cats -> sleep controlled at brain level Brainstem transect -> constant SWS, no REM sleep
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Describe the ascending arousal system in rodents
Different cell populations in brain that send projections inducing wakefulness. Lateral hypothalamus (LH) - orexin Tubermamillary nucleus (TMN) - histamine (anti-his pills make u feel drowsy), allergies Dorsal raphe (DR) - serotonin Locus coeruleus (LC)- oradrenaline ## Footnote Methamphetamines & ecstasy promote wakefulness by altering chemical levels
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Orexin (hypocretin)
Produced by neurons on lateral hypothalamus. - project across brain - signal through Oc1R & Ox2Rs in target regions ## Footnote Orexin KO show reduced wakefulness/increased SWS/REM -> promotes wake.
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Narcolepsy
Sleep disorder: - Excessive daytime sleepiness w/ irresistible sleep attacks - Cataplexy (episodes of paralysis/weakness preceded by strong emotions) - Sleep paralysis - Hypnagogic hallucinations (dream like images at sleep onset) ## Footnote 1 in 2000, appears 15-30yrs.
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What chemical is associated with narcolepsy?
Reduced orexin neurons found in post-mortem narcoleptic brains -> reduced orexin signalling. (also seen in dogs e.g. labradors)
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Unihemispheric sleep
Sleep w/ half of brain at a time - in aquatic animals & birds
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Sleep in mallard ducks
Can sleep w/ one eye open (1 hemisphere awake) When left eye closed (right hemisphere sleeping), vice versa -> decussation - Allows detection of predators during unihemispheric SWS (USWS) - Increase USWS use as risk predation increases - Bihemispheric sleep when predation risk is low ## Footnote Ducks on edge of row hgiher predation risk so more unihermispheric sleep
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Sleep in great frigate (Fregata minor)
Primarily rely on soaring-gliding strategy -> circular rising on thermal air currents (soaring). Used EEG to look at head position during wake & sleep. - During flight, SWS associated w/ tilted head - Sleep only occurs during ascending flight - Awake hemisphere is always opposite to direction of rotation (prevent collisions?) ## Footnote During flight, 95% time awake (only small amount of SWS) - most sleep during flight is ASWS and on land sleep is far more bihemispheric.
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Sleep in reptiles
Autralian dragon EEG & analysis of closed eyes -> regularly occurring episodes of REM, cycles take ~80s in this species
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How is sleep defined in smaller animals w/ simpler nervous systems?
- Period of quiescence associated w/ species specific posture - Increased arousal threshold - Quick reversibility to wakefulness - Homeostasis ## Footnote e.g. flies have preferred resting position + posture, have greater arousal threshold when resting + deprivation causes rebound rest (homeostatic)
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Sleep in insects
Described 1983 - now known in D. melanogaster, scorpions, honey bees & cockroaches. In honey bees: learning, memory, waggle-dance precision & navigational memory reduced by sleep deprivation. There is conservation of neurochemistry that control sleep-wake cycles between many species. -> drosophila can be used for sleep studies
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What are the two ways organisms adapt to cyclical changes in environmental conditions?
1. Passively/retroactively 2. Proactively
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What is the purpose of homeostatic mechanisms in organisms?
To regulate internal conditions at a set/optimal point in response to environmental changes
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What are biological timekeepers or clocks responsible for?
Controlling physiology and behaviour variations from day to night
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What are circadian rhythms?
24.5-hour cycles that are a ubiquitous control feature of physiology and behaviour
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List three physiological or behavioural changes regulated by circadian rhythms.
* Body temperature * Hormone secretion * Sleep
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What is necessary for an activity to be classified as a circadian rhythm?
It must be present in the absence of any external cues
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What does rodent wheel running activity demonstrate in constant conditions?
A near 24-hour free-running rhythm
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What happens when lights are turned on earlier by 6 hours in an experiment?
Activity slowly synchronises to the new cycle over multiple days
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True or False: Circadian rhythms are reactive to changes in environmental conditions.
False
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How do physiological and behavioural rhythms respond to environmental changes?
They anticipate changes, such as body temperature rising during the day and dropping before night.
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What type of activity do flies exhibit according to their circadian rhythms?
Crepuscular activity, being active at twilight
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What happens to male crickets' calling activity when kept in constant light?
They still show rhythmic calling, approximately every 26 hours.
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What is the role of the suprachiasmatic nucleus (SCN) in mammals?
It is the main mammalian circadian clock and acts as a pacemaker.
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What occurs after lesioning the SCN in animals?
Irregular bouts of activity, indicating the necessity of SCN for circadian rhythmicity
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What effect does SCN transplant have on lesioned animals?
It restores rhythmic behaviour.
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What happens to sleep rhythms in humans when the SCN is damaged?
Sleep rhythms become disrupted.
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What type of activity does the SCN exhibit daily?
Rhythmic activity in metabolic processes, with low activity at night.
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What was observed when individual SCN neurons were isolated and recorded?
They exhibited synchronous rhythmic activity when recorded in intact SCN tissue.
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What is the structure of the molecular clock in SCN cells?
A transcriptional/translational feedback loop comprising stimulatory elements (CLOCK/BMAL1) and inhibitory elements (PERIOD/CRYPTOCHROME) ## Footnote These elements refine the period of the clock cycle to 24 hours.
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Where was the molecular clock first discovered?
In cyanobacteria and plants ## Footnote Similar genes have evolved three distinct times.
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How is clock activity visualized in research?
Using transgenic mice that carry a bioluminescent reporter of clock gene activity ## Footnote Heterozygous PER2::LUC KI mice are used to observe diurnal expression of molecular clock genes.
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How does the SCN synchronize the biological clock to the environment?
SCN receives light information via the retinohypothalamic tract (RHT) ## Footnote The SCN is optimally positioned to receive retinal input from the optic chiasm.
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What happens to SCN cells in response to light?
Puts cells into a daytime state in vivo ## Footnote This is an example of acute adaptation.
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What mechanisms does the SCN use to communicate information to the rest of the body?
Coupling and effector mechanisms ## Footnote Increased neuronal signaling and endocrine signaling are involved.
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What role does melatonin play in circadian rhythms?
Regulates release from the Pineal Gland in response to light and circadian signals ## Footnote Melatonin is released at night in both nocturnal and diurnal animals.
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How does light affect melatonin release?
Light rapidly suppresses melatonin release ## Footnote SCN signaling inhibits production/secretion via a multi-synaptic SCN-Pineal pathway.
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What are the consequences of SCN lesions in wild chipmunks?
High mortality rate after 2 months compared to controls and sham-lesioned individuals ## Footnote This indicates the importance of SCN in survival.
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What happens when circadian rhythms are genetically manipulated in mice?
Reduced evolutionary fitness due to internal clock speed deviating from environmental rhythms ## Footnote This leads to a lower reproductive rate.
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What are ultradian rhythms?
Rhythms that operate over less than 24 hours ## Footnote Examples include human sleep cycles and pulsatile growth hormone release every 3 hours.
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What are infradian rhythms?
Long period rhythms greater than 28 hours ## Footnote Examples include bird migration and reproductive cycles.
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What is an example of a circannual rhythm?
Hibernation of the golden-mantled ground squirrel ## Footnote Hibernates from October to May, consistent without external cues.
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How do seasonal changes affect sheep physiology?
Length of coat increases in winter, size of testes and prolactin production increases in summer ## Footnote These changes are independent of external signals and regulated by melatonin rhythm.
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What controls prolactin release in relation to seasonal changes?
Pituitary pars tuberalis via melatonin rhythm ## Footnote This encodes day length and drives seasonal changes.
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Is the molecular clock basis fully understood?
No, it is not fully understood.
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What is the structure of the molecular clock in SCN cells?
A transcriptional/translational feedback loop comprising stimulatory elements (CLOCK/BMAL1) and inhibitory elements (PERIOD/CRYPTOCHROME) ## Footnote These elements refine the period of the clock cycle to 24 hours.
367
Where was the molecular clock first discovered?
In cyanobacteria and plants ## Footnote Similar genes have evolved three distinct times.
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How is clock activity visualized in research?
Using transgenic mice that carry a bioluminescent reporter of clock gene activity ## Footnote Heterozygous PER2::LUC KI mice are used to observe diurnal expression of molecular clock genes.
369
How does the SCN synchronize the biological clock to the environment?
SCN receives light information via the retinohypothalamic tract (RHT) ## Footnote The SCN is optimally positioned to receive retinal input from the optic chiasm.
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What happens to SCN cells in response to light?
Puts cells into a daytime state in vivo ## Footnote This is an example of acute adaptation.
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What mechanisms does the SCN use to communicate information to the rest of the body?
Coupling and effector mechanisms ## Footnote Increased neuronal signaling and endocrine signaling are involved.
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What role does melatonin play in circadian rhythms?
Regulates release from the Pineal Gland in response to light and circadian signals ## Footnote Melatonin is released at night in both nocturnal and diurnal animals.
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How does light affect melatonin release?
Light rapidly suppresses melatonin release ## Footnote SCN signaling inhibits production/secretion via a multi-synaptic SCN-Pineal pathway.
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What are the consequences of SCN lesions in wild chipmunks?
High mortality rate after 2 months compared to controls and sham-lesioned individuals ## Footnote This indicates the importance of SCN in survival.
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What happens when circadian rhythms are genetically manipulated in mice?
Reduced evolutionary fitness due to internal clock speed deviating from environmental rhythms ## Footnote This leads to a lower reproductive rate.
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What are ultradian rhythms?
Rhythms that operate over less than 24 hours ## Footnote Examples include human sleep cycles and pulsatile growth hormone release every 3 hours.
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What are infradian rhythms?
Long period rhythms greater than 28 hours ## Footnote Examples include bird migration and reproductive cycles.
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What is an example of a circannual rhythm?
Hibernation of the golden-mantled ground squirrel ## Footnote Hibernates from October to May, consistent without external cues.
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How do seasonal changes affect sheep physiology?
Length of coat increases in winter, size of testes and prolactin production increases in summer ## Footnote These changes are independent of external signals and regulated by melatonin rhythm.
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What controls prolactin release in relation to seasonal changes?
Pituitary pars tuberalis via melatonin rhythm ## Footnote This encodes day length and drives seasonal changes.
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Is the molecular clock basis fully understood?
No, it is not fully understood.
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Circatidal marine rhythms
Gravitational force of moon + rotational force of earth -> 12.4 tidal cycle. Moon overhead makes high tide higher (24.8 hrs cycle), exaggerated by moon/sun alignment (bi-lunar ~14 days).
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