Behavioral Biology - Behavioral Biology Book: animal behaviour  second or third international - Studeersnel (2024)

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• 22Notes, assignments and book summary of Behavioural BiologyBehavioural BiologyCollege-aantekeningen100%(1)
• 29Notes, assignment and book summary of Behavioural Biology for the 2nd examBehavioural BiologySamenvattingen100%(1)

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Behavioral Biology

• Book: animal behaviour  second or third international edition by oxford
• All chapters apart from 9

Lecture 1 (1-3)

Chapter 1 – the science of animal behaviour - What is an animal o An organism  Multicellular eukaryote  Dependent of organic materials  Breathe, move, nervous system, sensory system  Have blastula during early embryonic development!! - What is animal behaviour o Animal behaviour is internally coordinated externally visible pattern of activity that responds to changing internal or external stimuli  Internal: information processing  External: observable and measurable

 Also includes sleep and ignoring something anything an animal does

o It is now quantifiable  behaviour biologists quantify behaviour  Motor patterns  Different than just asking questions like psychologists  Measured with ethogram  Formal description of animal’s behaviour  Creation of a term like “feeding” and a very clearly defined description of the term (in your opinion)  Time budget: the total time of observation and relative frequency of behaviour

• Philosophy of behaviouro Proximate (how) Development Mechanismo Ultimate (why) Function Evolutiono Top-down vs bottom-up
• Avoid anthropomorphismo attributing human traits, emotions, or intentions to non-human entities
• Avoid anecdotal evidence
• Correlation and experimental evidence (cause and consequence)
• Systematic ((double) blind) observations
• Nature vs lab conditions
• Evolutionary context
• Interdisciplinary approachesChapter 2 – methods of studying animal behaviour
• Process of scienceo Knowledge/curiosity  question  preliminary observations  fine tuning of question ORformulate hypothesis  formulate specific predictions  determine the variables needed choose method of measurement  collect data  analysis  back to a new question Observational method

 An approach in which scientists observe an record the behaviour of an organism without manipulating the environment or the animal Causality is unclear Used to test hypotheses and describe behavioral patterns Experimental method An approach in which scientists manipulate or change a variable toexamine how it affects the behaviour of the animal Direction of causality is clear Comparative method An approach that examines differences and similarities between species tounderstand the evolution of behavioral traits Mainly for ultimate explanations Ancestral or derived traitso Ancestral: found in the common ancestor of two or more species pleiomorphic traito Derived: found in organisms but wasn’t present in last commonancestor  apomorphic trait Knowledge about relatedness Phylogeny  genetics Modelling Yields testable predictions Meta-analysis

• Ethics:o Scientific misconduct Violation of ethical behavior standard in scienceo The tree R’s

 Refinement  keep level of discomfort as low as possible

 Involves improving procedures and techniques to minimize pain and stress for animals

 Replacement  if another model is available (e. cell cultures), don’t do

experiment  Encourages the use of modeling, videotapes, or other approaches in place of actual animals

 Reduction  use minimum number of subjects to find an accurate enough average

 Promotes limiting the number of animals subject to disturbance in research

• Scientific knowledgeo Primary literature: the original source of scientific informationo Secondary literature: a report which often appears in newspapers, magazines, and books,that summarizes and interprets the primary literature
• Animal behaviour researcho Where to study animal behaviour Wild Semi natural conditions LabChapter 3 – evolution and the study of animal behaviour
• Evolution: a change in the inherited characteristics of biological populations over successivegenerations, 3 conditions are needed:o Heredity (DNA/RNA, genes) Testing whether a trait is heritable Parent-offspring regressiono Statistical technique used to examine the similarity betweenparents and their offspring in terms of the traits they posses Selection experimento Experiment in which different groups of individuals are subject todifferential selection on a trait

 Direct fitness: genes contribute to the next generation by an individual due to its own reproduction Indirect fitness: the genes contributed to the next generation by an individual as a result of helping non-offspring kin produce additional offspring Inclusive fitness: the sum of individuals direct and indirect fitness Kin selection: a form of natural selection in which individuals can increase their fitness by heling close relatives because close relatives share genes Multilevel selection: a form of natural selection that may result in circumstances when selection on groups is stronger that selection on individuals Individuals have a genotype and phenotype; however, selection only acts on the phenotype as the DNA needs an environment to develop and act out a phenotype that can be selected for  Phenotypic plasticity  genotype can develop into numerous phenotypes Maternal and transgenerational effects (non-genetic inheritance)  For the individual  genotype x environment  Gene-environment interactions shape phenotypes

Lecture 2 (5-6)

Chapter 5 – sensory systems - The environment acts on the sensory systems which result in sensory processing in the brain as well as behavioral output o Environment  abiotic and biotic factors  Temperature  Light intensity  Food  Mates  Predators  Competitors o Sensory processing  Transformation of the sensory input into information  Produces a (behavioral) response o Behavioral output  Mating behavior  Individual recognition  Orientation  Prey detection  Predator avoidance o The interactions of the environment and the sensory systems are what result in fitness and eventually evolution - Sensory structures gather energy from the environment  channel it to specialized nerves  produce an electrical signal in the nervous system o Environmental energy  Light  Heat  Sound  Electricity  Gravity  Magnetism  Chemical bonding - Chemoreception o The oldest and most widespread sensory system o Taste: gustatory receptors detect dissolved molecules o Smell: olfactory receptors detect dissolved molecules - Photoreception

o Light detectiono Image formationo Colour discriminationo Photoreceptors: cones and rods  Cones: colour  Rods: low light intensity visiono Differences amongst species

• Mechanoreceptiono Detection of vibrations (i. mechanical waves) that travel through air, water or othersubstanceso Mechanoreceptors are present on the head, thorax, wings, legs, etc.o Sound as mechanoreception Species differ greatly in the frequencies of wavelengths they can detecto Lateral line system in fish Detects changes in water pressure created by other individuals (movement in thewater) An attacking predator produces a flow stimulus  prey lateral line receptors detectthe predator flow  the prey executes a fast starto Substrate-borne vibrations Many arthropods communicate through vibrations transmitted through solidobjects
• Electroreceptiono Detection of electrical fields
• Magnetoreceptiono Detection of magnetic fieldso Physiological mechanism largely unknowno Receptors not identifiedo May be present in several animal taxa
• Predator prey co-evolutiono Interaction between species  fitness  evolutiono Predator prey sensory systems co-evolve as well Many insects can detect 2-100 kHz frequencies, corresponding to the typicalecholocation frequencies of bats Echolocating bats vary the rate of emitted ultrasonic pulses Ability evolved at least 18 times independently Selection can favor predator sensory systems that enhance their ability to detectprey Selection can also favor prey that possess adaptations that reduce the foragingabilities of their predatorsChapter 6 – communication
• Communication  a specialized signal from one individual that influences the behavior of anotherindividualo Signaler: information  encode  signalo Receiver: perceive  process  response
• Signalo Evolved train that can take many forms Vocalizations Chemical pheromones Morphological traits Electrical discharges Substrate borne vibrations Behavioral displayso Signal versus cues Signals: Any act or structure which alters the behaviour of other organisms Have evolved because their effects benefit the signaler

o signals will be accurate indicators when they are costly to produce or maintain when signal senders can benefit from dishonesty, signal receivers will beselected to resist manipulationo co-evolutionary arms raceo Environmental factors Environment influences the evolution of signals  different environments requiredifferent signals Chemical signalso Long lasting and travel great distanceso Not modified once transmitter Auditory signalso Travel around objectso Turn on/off quicklyo Diminish with distance Visual signalso Move rapidlyo Require adequate light levels Color signal effectiveness was affected by light environment, for exampleon brightly colored fish who use their colors to attract mates. The deeperthey are the less vibrancy to their colors Habitat structure favors vocal signals that minimize attenuation Urban sounds affect signal production Extended phenotypes Expressed beyond the body of an individual Often involve modification of the environment Costly to build and maintain Signal interception by others Bystanderso A third party individual that detects a signal transmitted betweena signaler and a receiver Audience effecto Occurs when the presence of a bystander influence the behaviorof the signaler

Lecture 3 (4)

Chapter 4 – behavioral genetics - Individual differences in behaviour o Genetic and environmental influences  Phenotype – variation in observable traits  Behavior  Morphology  Physiology  Genotype – variation due to differences in genetic makeup of the organism  Do genes encode for behavior o Genes code for proteins, not for behavior and the road from proteins to behavior is long  DNA  RNA  proteins  brain/physiology  behavior  Why does it matter  The origin of individual differences determines how behavior may change o Only when part of the phenotypic differences has a genetic basis, these traits can evolve

 When these differences affect fitness, natural selection may lead to adaptive evolution of behavioro Only when there are environmental influences, can behavior bemodulated Adaptive evolution of behavior Evolution can shape behavior through natural selection, when (at least partof) its variation is heritable and it affects:o Phenotypic variation among individuals which has genetic basiso Differences among individuals/genotypes in survival/viability andreproductive successo Consistent differences in survival and reproduction, associatedwith the same genotypes (natural selection)o Change in allele frequencies in the population (evolution) Earliest evidence of a genetic basis Instinct or innate behavioro Behaviors that are performed the same way each time fullyexpressed the first time they are exhibited, and present even inindividuals raised in isolation Reflexeso Involuntary movement in response to stimulus Fixed action patterno Behavior that displays almost no variation and once startedcannot be stopped until completed Environmental influences on behavior Environment during early developmento Prenatal conditionso Imprinting and habituationo Social environmento Experience and learningo Food/diet and stresso More Current environment individual variation caused by environment is not inheritedo Plasticity and genetic + environmental interactions Phenotypic plasticity Changes in phenotype/ behavior in response to environmental conditions Reaction norm The range of behaviors expressed by a single genotype in differentenvironments The same genotype across different environmental conditions results inrange of behavioral phenotypeso Genotypes may exhibit similar response to environmento Genotypes may respond differently to environment resulting ingene environment interaction Gene environment interaction When environment has greater effect on one genotype that the others Most likely the rule rather than exception in behavior Natural selection can alter the plasticity of traits when there are gene +environment interactionso Individual personalities Consistent or repeatable inter-individual differences in behavior over time or acrossenvironmental contexts  personality Attributable to the combined influences of genetic effects andenvironmental effects May place constraints on individual variation Behavioral syndromes

o Individual gene that is responsible for large fraction of phenotypic variationo E. rover and sitter foraging behavior in fruit flies  Observation:  Drosophila melanogaster larvae exhibit two behavioral polymorphisms  Rovers have longer foraging trials in presence of food and are more likely to leave a food patch than sitters  Polymorphism maintained by frequency dependent selection  Rover has higher fitness in crowded environment  Sitter has higher fitness in less crowded environmento QTL analysis: to detect major effect loci  Find loci that contributes to the phenotypic variation  Statistically associate phenotypic trait variation with allele at detectable genetic markers  Associates genetic info with trait info  QTL are stretches of SNA that either contain or are linked to genes influencing a trait such as behavior  QTL mapping is a statistical technique that combines genetic information with trait information to determine which regions of the genome contain the genes that influence the trait QTLs  Candidate genes are major genes suspected of contributing to a large amount of the phenotypic variation in a Specific trait  Genetic markers  detectable mutations that geneticists use to recognize specific regions of chromosomes  Not assumed to be causal alleles but linked to  Higher number of markers means narrower regions can be identified as containing a putative GTL  Main requirement of a marker: polymorphic in the population under study  Analysis  Select two strains that differ in a quantitative trait  Cross + back cross: recombination generates a mosaic of the 2 parental chromosomes  Score each individual for the phenotypic trait  Determine genotype of each individual for each marker  Statistics: which markers are associated to the phenotypic trait  Limitations  The number of loci that contribute to the variation is less than those that contribute to the development of a trait o Requires crossing the different strains/species

o Only loci that are variable int eh populations under study can be detectedo Especially major effect genes can be detected Minor gene:o Individual gene that contributes to small amount of variation inthe phenotype  may be rule rather than exceptiono Minor effect loci difficult to detect with most methods  onlyminor contributions to the trait variation

Lecture 4 (7)

Chapter 7 – Learning - Definitions of learning o That process which manifests itself by adaptive changes in individual behaviour caused by experience o Takes place when some experience results in a relatively permanent change in the reaction to the situation - Learning is a form of phenotypic plasticity o The phenomenon that a genome can result in different phenotypes - Learning curves o How much an animal learns and therefore the shape of the curve is determined by the difference between what an animal already knows and what it perceives. But also of the learning speed which is dependent on the relationship between natural behaviour of an animal and the specific learning task. Hence the delta rule:  A basal form of the delta rule; this rule can be much more complicated - When, what, how much and why to learn is dependent on the costs and benefits, and is therefore related to the increase in fitness o Two factors affect the evolution of learning  Environmental stability  The usefulness of past experience o when will learning evolve  when there are new things to learn  like in a changing habitat or when past experience isn’t sufficient to maintain fitness in a current state - forms of individual learning o non-associative learning  uses a single stimulus to activate learning  sensitization  repeated administration of a stimulus results In the progressive amplification of a response  habituation  innate, non-reinforced response to a stimulus decreases after repeated or prolonged presentations of that stimulus o associative learning  uses stimulus-stimulus activation apart from classical conditioning (uses response – reinforcement activation)  classic conditioning  neutral stimulus becomes associated with a stimulus that naturally produces a behavior  operant (instrumental) conditioning

o to master this, it is essential to read books and literature

• animals find food by integrating sensory modalitieso sensory modalities vision olfaction taste touch auditory vestibular proprioception electric infraredo integrating senses improves foraging and robustness
• visual predators find cryptic prey by learning a search imageo prey can be located using the search imageo the prey is identified using eye, head shape and hind leg shape identificationo some animals use find cryptic prey by learning a search image this reduces search time
• optimal diet predicts how animals should include food typeso optimal diet is based on optimal foraging theory optimal foraging theory assumes: natural selection have favored feeding behaviors that maximize fitness fitness while feeding increases with energy intake rate optimal diet model assumes: foragers maximize fitness by maximizing energy intake rate food items are encountered one at a time in proportion to their abundance food items can be ranked by their profitability
• optimal patch use predicts how food patches should be exploitedo the optimal patch use model assumes foragers attempt to maximize energy intake rate all patches are identical travel time between patches is constant the instantaneous harvest rate declines as a forager depletes a patch the forager experiences diminishing returns in each patcho optimal patch use time depends on travel timeo food intake give-up density depends on fitness risk many nocturnal animals perceive a higher predation risk at high light levels humans add artificial light to many environments which can negatively affect rarespecieso incomplete information and food patch estimation optimal patch use model assumes animals know the quality of encountered patches however, many foragers will need to estimate food patch quality animals can combine sample information from a patch with prior knowledge aboutthe distribution of patch types in the environment using Bayesian estimation combine prior observed intake statistics for patches to correct patchquality estimate based on current intake
• in groups, some conspecifics take food discovered by otherso within a social foraging species we often see food producers and scroungers producers: search for food scroungers: usurp food discoveries from producerso producer scrounger model predicts the frequency of each forging strategy in a groupo model predicts variation in the frequency of scroungers such that there is equal fitness forboth strategies within a groupo this trade off is also observed in behavioral manipulations
• foraging underpins animal training and welfare

o training works for all animals from invertebrates to vertebrates because all forage and learn based of habituation and operant conditioningo animal training is useful to improve welfare or pets and their interactions with humanso animal training is useful to improve welfare of farm animals from fish to birds to horseso animal training is useful to improve welfare of animals generally also in zoos and parkso animal training process from breeding to research  design operant conditioning training process based on ecology and foraging behavior  starting early, without taking animals away from parents. This maximizes welfare  the same process applies for animals kept in zoos, parks, farms, and as petso train one behavior at a time and split behavior up in stepso chaining: animals combine individual trained behaviors  order doesn’t matter as foraging behaviors are generalized

Lecture 6 (11)

Chapter 11

Lecture 7

Ontogeny - ontogeny/ ontogenesis o general term for development o formation of a being o this term is not only used in a behavioral context o not only about proximate questions  can we better understand the organization of complex adult behaviour when we follow its development  flexibility and underlying mechanisms o also about function and evolution o nature versus nurture:  innate/instinct/inherited vs learning/experience  tabula rasa  deprivation studies used to research the influences of both  song in chaffinch birds o reared with hearing others: normal song o reared in isolation: abnormal song o reared in isolation and deaf: abnormal song  deprivation leads to alterations in the visual system  Harlow: studied the mechanisms by which newborns rhesus monkeys’ bond with their mothers  More time with furry, more exploration (less fear) with furry  Conclusion: sensitive response and security more important than food for attachment  Animals became very poor mothers themselves o Imprinting  A phase sensitive rapid form of learning (independent of the consequences) about a certain stimulus in a critical period  A life stage in which the nervous system is especially sensitive to certain stimuli

 An individuals biological sex can be genetically or environmentally determined or by a combination of both depending on species An individual may even be both male and female or may switch sex Sex differentiation in birds Estrogen turns males into females Blocking estrogen synthesis turns females into males Birds: E2 feminizeso Mammals: T masculinization A sex difference in the songbird braino Exposing females to T or E2 gives male like vocal control regions organizing effecto Exposing these females later to T: sing like male  activatingeffect Sex Large variability in systems to become male or female Important because reproductive investment/behaviour Variability in behaviour within sexes depending on the rearing environmento play behaviour performed in the absence of normal stimuli or behavior elicited by normal stimulibut not followed to the completion of the ritualized behavior pattern. Play has beendocumented only in mammals and birds common among immature animals, apparently part of the process of learning adultbehavior much of play in mammals serves to develop hunting skills adult animals also engage in play horses, cattle, and other hooved mammals run, chase, kick up for noobvious reason dogs have postural signals of mock aggression used to entice others intoplay all elements of ritualized behavior may be present but do not follow pattern orsequence necessary to communicate serious intent 5 criteria for play Intrinsically rewarding Does not seem to have an obvious purpose Repetitive Voluntary Under relaxed conditions  healthy/fit animalso Lambs decrease play behavior under bad food conditionso Kittens increase play in bad food conditions Why play Development of motor control Forming social bonds Establishing a social hierarchy Learning species specific/sex specific adult behaviors Learning about the environment Fun  pleasant Types of play Locomotor play Object play Social play Who plays Rough and tumble  and pursuit play in rhesus monkeys Decrease in play into adulthood Effect of androgens Sex differences Males and females play differently, also in their choice of objects

 Human play  The ultimate players  Play amongst each other  A lot with other species  Dogs also initiate playing with people Conclusion  Multiple types of play  Multiple reasons why animals play  Sex differences  Decrease with increasing age  Neglected field in behavioral bioo parental and environmental effects when the phenotypes of the parents influence the phenotype of their offspring,these are called parental effects feather pecking poultry behavior industry and amateur husbandry  not observed in wild abnormal/undesirable cutting beak: other problems possibly problems during development psychopathology early feather pecking:o gentle feather peckingo present directly after hatchingo possibly a form of social exploration ground pecking: foraging dustbathing social recognition important for forming stable hierarchies social recognition is a lateralized trait lateralization is affected by light during incubation, physiologically at leastduring the first three weeks after hatching conclusion there is no reason to believe that even though the presence of substrate isextremely important in preventing development of feather pecking there are social factors that play a role and which are affected byenvironmental factors severe feather pecking seems to rapidly develop from gentle even though animals may have strong predispositions this is not necessarilygenetic, environmental factors shape their phenotype these environmental factors may include self, playing increasesexperiences which may be individual specific environments in which animals have not evolved may yield quite differentphenotypes and may lead to behavioral abnormalities

• conclusiono ontogeny is about proximate and ultimate questionso ontogeny is the continuous translation interaction between G and Eo the nature vs nurture debate is about a wrong dichotomyo sensitive periods with long lasting consequences sexual differentiation imprinting

Lecture 8 (15-16)

 resource defense  clownfish live in social groups that defend a sea anemone. They eat undigested food from the anemone and use it for a breeding site  the larger the anemone the more effort expended to defend it social learning  intelligent animals can learn about foraging or other opportunities from other animals in their social group cooperation/division of labor  behavior that is mutually beneficial for the fitness of both parties in the behavior  giving an alarm group increases predation risk for caller, but lowers for other members of the group  there can therefore still be a net fitness benefit from an emergency call by Kin selection o phenomenon that fitness also depends on breeding success of relatives  hamiltons rules: carry out an altruistic act when: o rb – c > 0  r = coefficient of relatedness between actor and recipient  b = direct benefit to recipient  c = direct cost to actor  rb = indirect benefit to actor  measured in terms of evolutionary fitness  kin selection in ground squirrels o sons disperse more than daughters o females are more related to group than males o prediction: females make more alarm calls than males because the indirect benefit is larger for females  kin discrimination o for kin selection to operate, individuals must be able to discriminate between kin and non-kin o done through:  direct familiarization  learned familiarity  specific individual recognized based on previous experience of those individuals  indirect familiarization  phenotype matching  close relatives may have similar smell, eye color, hair color, etc.  spatial recognition  in species with low dispersal, individuals living closest to you are most related  cooperative breeding o in some social groups, a few adults reproduce while others help raise their offspring o kinship favors cooperative reproduction. Helpers are often closely related to the breeders, usually older children from previous breeding bout. Hence selection will favor cooperation o ecological constraints favor cooperative reproduction  lack of breeding sites will increase the cost of independent breeding  social queueing inheritance of breeding site  cooperative behaviour amongst unrelated individuals o byproduct mutualism  each individual behaves in a selfish way but when they act together there is a side effect of additional benefit to cooperators

o reciprocal altruism  individuals act altruistically when there will be repayment of the altruistic act at a future time

• parental care in behavior that increases the fitness of offspringo high levels of parental care are typically found in species that produce small numbers of largeoffspringo parental care varies across species and reflects life history trade offs in life histories a tradeoff is the causal negative relationship between two fitnesscomponents energy and time (effort) are limited  effort allocated to reproduction reduceseffort that can be allocated to an individual’s growth and survivorship relevant traits include age at first reproduction, number and size or offspring, levelof parental care, mortality rateo the example of the tree hopper research question: within a species, why is there much variation in parental caresuch as egg guarding hypothesis: there is tradeoff between current and future reproduction prediction 1: increased egg guarding will increase egg survival (currentreproduction) prediction 2: increased egg guarding will decrease future reproduction conclusion: egg guarding enhances current reproduction at the expense of futurereproductive success.
• Sexual conflict and parental care:o internal versus external fertilization Birds and mammals have internal fertilization  it is easy for a male to abandonoffspring but difficult for the mother to do so  more common to have maternalcare Many fish have external fertilization  it is now easy for both males and females toabandon offspring  more common to have minimal parental careo Determinate versus indeterminate growth Birds and mammals have determinate growth  once a female reaches adult size,energy can be devoted to reproduction rather than additional growth  increasedparental investment Many fish have indeterminate growth  females must trade off currentreproduction versus future growth  low investment of maternal careo Certainty of paternity With internal fertilization, male birds and mammals may have little confidence thatthey are actually the father. Parental care is expensive because fathers may end upinvesting in offspring that is not their own  maternal care more often With external fertilization, males have much more confidence of paternity evolution of mate guarding  male parental care more commono Parent-offspring conflict The fitness benefit for offspring of receiving parental care is twice the fitness for anadult of providing care, because the coefficient of relatedness between parent andoffspring is 0. Optimal strategy for the mother: provide equal amount of investment to eachoffspring Optimal strategy for the offspring: take more investment than siblings More related to yourself than your siblings You can’t be certain that your sibling has same father so relatedness maybe low lower level of relatedness between siblings will increase the severity of parentoffspring conflict brood reduction