ANSC3090 Animal Behaviour Lecture 10-19 PDF

ANSC3090 Oct 8, 2024 Picture: Dr. Kutzer Alexandra Harlander Dr. vet. med. PhD, Dip ECAWBM(WSEL) 1 Video: Dr. Reuckli 2  400,000...

ANSC3090 Oct 8, 2024 Picture: Dr. Kutzer Alexandra Harlander Dr. vet. med. PhD, Dip ECAWBM(WSEL) 1 Video: Dr. Reuckli 2  400,000 346,791 350,000 310,961 300,000 250,000 # hits 200,000 150,000 140,831 100,000 55,286 50,000 14,823 7 107 385 5,039 0 1950 1960 1970 1980 1990 2000 2010 2020 Decade 3 4 Behaviour: What do we mean by this word? Can only animals behave, or can any living organism exhibit behaviour? Resting outside Is inactivity considered a behaviour? Can groups behave, or is behaviour limited to individuals? Must behaviour involve motion? Can developmental changes in response to a stimulus be Foraging inside considered behaviour? 5  Behaviour: Definitions? Do plants behave? Plants grow towards or away from stimuli. Plants release chemicals in response to diseases. Plants respond to changes in season. Plants adapt and survive in their environments. Definitions are crucial. Resting outside Definitions of terms reflect the current knowledge and understanding at a specific time. 300-year-old lime tree Ethograms: practical need to define behaviour Foraging inside 6 Examples 7 Definitions of Behaviour (dictionaries) ‘the way that someone behaves’ ‘the way that someone or something behaves in a particular situation’ ‘the way a person or animal acts or behaves’ ‘the way something (such as a machine or substance) moves, functions, or reacts 8 Definitions of Behaviour (Behavioural Biology) “The total movements made by the intact animal.” Tinbergen (1951) 9 Definitions of Behaviour (Behavioural Biology) “What an animal (or plant) does.” Davis (1966) 10  Definitions of Behaviour (Behavioural Biology) “Behaviour includes all those processes by which an animal senses the external world and the internal state of its body and responds to changes it perceives.” Manning (1972) 11 Definitions of Behaviour (Behavioural Biology) “Behaviour can be defined as the way an organism responds to a stimulation.” Raven and Johnson (1989) 12 Definitions of Behaviour (Behavioural Biology) “External visible activity of an animal, in which a coordinated pattern of sensory, motor, and associated neural activity responds to changing external or internal conditions.” Beck et al. (1991) 13 Definitions of Behaviour (Behavioural Biology) “Observable activity of an organism; anything an organism does that involves action and/or response to stimulation.” Wallace et al. (1991) 14 Definitions of Behaviour (Behavioural Biology) “A response to external and internal stimuli, following integration of sensory, neural, endocrine, and effector components. Behaviour has a genetic basis, hence is subject to natural selection, and it commonly can be modified through experience.” Starr and Taggart (1992) 15 Definitions of Behaviour (Behavioural Biology) “All observable or otherwise measurable muscular and secretory responses (or lack thereof in some cases) and related phenomena such as changes in blood flow and surface pigments in response to changes in an animal’s internal and external environment.” Grier and Burk (1992) 16 Definitions of Behaviour (Behavioural Biology) “Behaviour involves the interaction between an animal’s machinery, its bones, muscles, nervous system, etc. and its outside world, such as its food, enemies and social practice.” Hall and Halliday (1998) 17 Definitions of Behaviour The challenge 18 Definitions of Behaviour Survey study of 174 respondents with varying levels of knowledge about behavioural biology. Asked to indicate whether a given phenomena is a behaviour or not. Asked to agree or disagree with various statements about behaviour. 19 Definitions of Behaviour Generally approved (≥64% agreement) statements: A developmental change is usually not a behaviour Behaviour is always influenced by the internal processes of the individual Behaviour is something whole individuals do, not organs or parts that make up individuals A behaviour is always in response to a stimulus or set of stimuli, but the stimulus can be either internal or external 20 Definitions of Behaviour Generally disapproved (≤24% agreement) statements: Behaviour is always in response to the external environment A behaviour is always an action, rather than a lack of action People can all tell what is and isn’t behaviour, just by looking at it Behaviour always involves movement Behaviours are always the actions of individuals, not groups In humans, anything that is not under conscious control is not behaviour Behaviour is always executed through muscular activity 21 In summary No one can agree on how behaviour is defined (more than 50% gave contradictory answers) Why does this matter? Makes it difficult to tackle significant moral/emotional problems Intellectual rigour of scientific inquiry 22 The result? “Behaviour is the internally coordinated responses (actions or inactions) of whole living organisms (individuals or groups) to internal and/or external stimuli, excluding responses more easily understood as developmental changes.” Levitis, Lidicker Jr., & Freund (2009) 23 Criteria of definitions 1. Operational – allows the inclusion or exclusion of boundary phenomena 2. Essential – it is based on what survey responses intuit the word to mean 3. Widely applicable – avoids taxonomic bias and can be applied to any living organism 4. Succinct – each word is necessary 24 Terminology – e.g. Dictionary of Animal Behaviour Access through the University Library 25 Context and levels of behaviour/behavioural analysis Macro – Whole animal Individual – Groups of Laboratory – Field – Highly controlled Natural conditions Species – Population of Micro – (Edwards, 2018) Cellular 26 Context and levels of behaviour/behavioural analysis Species Seasonal timing of migration Behavioural type Feeding Population Migration Behavioural act Biting grass blades Family group Flying to food source Body parts Leg Dyad Agonistic interaction between Muscles Gastrocnemius individuals Individual Walking to food patch Neurons Tibial nerve (Edwards, 2018) 27 https://mediahelpingmedia.org/2021/02/04/the-questions-every-journalist-should-ask/ (Tinbergen, 1963) 28 Four Questions and Proximate and Ultimate Explanations Proximate Factors – HOW? Directly trigger and control behaviour e.g., physiological, developmental proximate causes 29 Four Questions and Proximate and Ultimate Explanations Ultimate Factors – WHY? Related to consequences or adaptive value of a specific behaviour 30 Proximate and Ultimate Explanations Example: Foraging Behaviour Video: Dr. Reuckli 31 Tinbergen’s ‘4 Questions’ Why do animals behave the way that they do? (‘how’ and ‘why’) Four ‘levels of explanation’ – Equally important to understand animal behaviour Framework for thinking about animal behaviour 32 (Tinbergen, 1963; Bateson and Laland, 2013; Beer, 2019) Tinbergen’s ‘4 Questions’ Causation Survival value Animal behaviour Tinbergen’s 4 Questions Ontogeny Evolution (Tinbergen, 1963; Bateson and Laland, 2013; Beer, 2019) 33 Tinbergen’s ‘4 Questions’ Current/One point in time Causation Survival value Mechanistic (Adaptive) Function (Current) Utility Animal behaviour Tinbergen’s 4 Questions Ontogeny Evolution Developmental Phylogenetic Chronological/Historical (Tinbergen, 1963; Bateson and Laland, 2013; Beer, 2019; 34 Dugatkin, 2019; Nesse 2013)  Tinbergen’s ‘4 Questions’ Current/One point in time Causation Survival value Mechanistic (Adaptive) Function (Current) Utility Proximate Animal behaviour Ultimate Questions/ Tinbergen’s 4 Questions Questions/ Explanations Explanations (‘how’) (‘why’) Ontogeny Evolution Developmental Phylogenetic Chronological/Historical (Tinbergen, 1963; Bateson and Laland, 2013; Beer, 2019; 35 Dugatkin, 2019; Nesse 2013) When measuring animal behaviour…. Confident? Happy smile? On January 31, 1961 Ham became the first chimpanzee in space. Clever Hans, phenomenon: a performing horse in Berlin in the late 19th and early 20th centuries – did Hans solve calculations? 36 Folk psychology ? Folk psychology part of our daily life? Human psychology is founded on folk psychology: e.g. natural interpretation of children’s behaviour given our robust knowledge of human children. Folk animal psychology? The kind of expertise that humans have when they spend a lot of time interacting with another species. Farmers, zookeepers, vets, pet owners develop a folk animal psychology that they use to understand, predict and better interact with the animals in their care (Andrews 2009, 2015) 37 Anthropomorphism? The attribution of human characteristics and feelings to other animals Starting point for considering what animals want/need in order to have a good life? It is essential to use science and other information when considering/evaluating animal welfare, not relying on anthropomorphism alone. 38 Critical Anthropomorphism Critical anthropomorphism Essential not to rely on anthropomorphism alone, without science [SCIENTIFIC DESCRIPTION] or other information 39 Structured approaches Observing to observing behaviour animal behaviour involve defining ethograms and describing the Single observed behaviour animal using time parameters 40  Observing animal Behavioural observations behaviour Group of animals Scientific description of behaviour Structured approach 41 Behavioural observations – a structured approach 1) Which animals to observe? 2) What behaviour to record? It all depends on your 3) Define behaviour (ethogram) research 4) Data collection protocol (sampling question/hypothesis! See one of the next lectures methods) 5) When should the animal(s) be observed? 6) Preliminary observations, pilot study 42 Which of these cats are ‘lying down’? Photo by Jacalyn Beales on Unsplash Photo by Alvan Nee on Unsplash 43  Which of these cats are ‘lazy’? Photo by Jacalyn Beales on Unsplash Photo by Alvan Nee on Unsplash 44 Which of these cats are ‘lying sternally’? Photo by Jacalyn Beales on Unsplash Photo by Alvan Nee on Unsplash 45 Behaviour measurements Quantitative Qualitative - How often? - Whole animal - How long? approach - How quickly? - Descriptive terms - Etc. - Qualitative Behavioural Assessment (QBA) (Travnik and Sant’Anna, 2021) 46 Qualitative Behavioural Assessment (QBA) Photo by Jacalyn Beales on Unsplash Not relaxed Completely relaxed (Travnik and Sant’Anna, 2021) 47 Behaviour measurements Quantitative Qualitative - How often? - Whole animal - How long? approach - How quickly? - Descriptive terms - Etc. - Qualitative Behavioural Assessment (QBA) (Travnik and Sant’Anna, 2021) 48 Ethogram ‘a catalogue of descriptions of the discrete, species typical behaviour patterns that form the basic behavioural repertoire of the species’(Martin & Bateson, 2007; p.34) 49 Ethogram: directories/catalogs of species-specific behaviours A list of behaviours with clear definitions/detailed descriptions Mutually exclusive Specific to the animal you are studying Can be simple or very complex Should include ‘” other” and /or “out of view” Objectively described in detail so that another observer can recognize the behaviour. It is important to test it out and practice! 50 Example Ethogram 51 Example Ethogram 52 Terminology – Time-parameters of behaviour Duration - State Frequency – Event Interval Pause Latency (Altmann, 1974) 53 Terminology- Duration States: have a duration, long (e.g., seconds, minutes, hours, % of time) Should be mutually exclusive Example: DURATION: 450 sec = 7.5 minutes = 75% feeding within the 10 minutes observation period (Altmann, 1974) 54 Terminology - Duration Pauses between behavioural states (Altmann, 1974) 55 Terminology – Frequencies and Rates Events Frequency: counts Rates: events/number of occurrences per unit time; distribution over time Example: RATE: 3x/600sec = 0.005 (Altmann, 1974) 56 Terminology – Latency Latency: duration (seconds, minutes, etc.) Time it takes to perform a specific behavioural pattern Example: (Altmann, 1974) 57 Terminology – Behavioural Bouts Bout: sequence of actions that are temporally clustered together Example: Repetitive pecking followed by a pause 58  leI ANSC3090 Oct 10, 2024 Picture: Dr. Kutzer Alexandra Harlander Dr. vet. med. PhD, Dip ECAWBM(WSEL) 1 Structured approaches Observing to observing behaviour animal behaviour involve defining ethograms and describing the Single observed behaviour animal using time parameters 2  Observing animal Behavioural observations behaviour Group of animals Scientific description of behaviour Structured approach 3 Behavioural observations – a structured approach 1) Which animals to observe? 2) What behaviour to record? It all depends on your 3) Define behaviour (ethogram) research 4) Data collection protocol (sampling question/hypothesis! See one of the next lectures methods) 5) When should the animal(s) be observed? 6) Preliminary observations, pilot study 4 Which of these cats are ‘lying down’? Photo by Jacalyn Beales on Unsplash Photo by Alvan Nee on Unsplash 5  Which of these cats are ‘lazy’? Photo by Jacalyn Beales on Unsplash Photo by Alvan Nee on Unsplash 6 Which of these cats are ‘lying sternally’? Photo by Jacalyn Beales on Unsplash Photo by Alvan Nee on Unsplash 7 Behaviour measurements Quantitative Qualitative - How often? - Whole animal - How long? approach - How quickly? - Descriptive terms - Etc. - Qualitative Behavioural Assessment (QBA) (Travnik and Sant’Anna, 2021) 8 Qualitative Behavioural Assessment (QBA) Photo by Jacalyn Beales on Unsplash Not relaxed Completely relaxed (Travnik and Sant’Anna, 2021) 9 Behaviour measurements Quantitative Qualitative - How often? - Whole animal - How long? approach - How quickly? - Descriptive terms - Etc. - Qualitative Behavioural Assessment (QBA) (Travnik and Sant’Anna, 2021) 10 Ethogram ‘a catalogue of descriptions of the discrete, species typical behaviour patterns that form the basic behavioural repertoire of the species’(Martin & Bateson, 2007; p.34) 11 Ethogram: directories/catalogs of species-specific behaviours A list of behaviours with clear definitions/detailed descriptions Mutually exclusive Specific to the animal you are studying Can be simple or very complex Should include ‘” other” and /or “out of view” Objectively described in detail so that another observer can recognize the behaviour. It is important to test it out and practice! 12 Example Ethogram 13 Example Ethogram 14 How do you design an ethogram? 100h of observations Literature review Visually Reviewed by 18 experts representing the behaviour 38 play behaviours - Locomotor play (14) - Object play (14) - Sexual play (3) - Play fighting (7) 15 How do you design an ethogram? Challenges: start and end of a behaviour Overlapping behaviours Complex behaviours Observer bias Solutions: Clear definitions, training, use of technology, context 16 Once you have defined the behaviour in an ethogram…. 17 Terminology – Time-parameters of behaviour Duration - State Frequency – Event Interval Pause Latency (Altmann, 1974) 18 Terminology- Duration States: have a duration, long (e.g., seconds, minutes, hours, % of time) Should be mutually exclusive Example: DURATION: 450 sec = 7.5 minutes = 75% feeding within the 10 minutes observation period (Altmann, 1974) 19 Terminology - Duration Pauses between behavioural states (Altmann, 1974) 20 Terminology – Duration Define time units & be consistent Record duration from start to end 21 Software designed for analyzing audio/video/ life observations (examples) www.noldus.com www.mangold-international.com 22 Terminology – Frequencies and Rates Events Frequency: counts Rates: events/number of occurrences per unit time; distribution over time Example: RATE: 3x/600sec = 0.005 (Altmann, 1974) 23 Terminology – Latency Latency: duration (seconds, minutes, etc.) Time it takes to perform a specific behavioural pattern Example: (Altmann, 1974) 24 Terminology – Behavioural Bouts Bout: sequence of actions that are temporally clustered together Example: Repetitive pecking followed by a pause 25 Terminology actor/recipient Social interactions Actor: animal that initiates the behaviour Recipient: animal that receives/is subject of the behaviour Need to ID animals? Depends on your research question! 26 Other considerations Two important ‘behaviours’ - Other Especially for time budgets - Out of sight Avoid ambiguity/being vague (Altmann, 1974) 27 Recording rules/methods 28 Rules for data recording & sampling Sampling rules (who, what) Ad lib Focal Scan Behaviour sampling sampling sampling sampling Recording rules (when) Continuous Time recording sampling = Interval structured recording Instantaneous One-zero sampling sampling 29 (Martin and Bateson, 1986, Altmann, 1974) Rules for data recording Recording rules (when) Continuous Time recording sampling For a specific amount of time You can get true frequencies, latencies, Instantaneous One-zero durations, etc. sampling sampling 30 (Martin and Bateson, 1986, Altmann, 1974) Continuous recording 31 Rules for data recording Recording rules (when) Continuous Time = Interval structured recording recording sampling At pre-determined intervals (every 30 sec, every 5 min etc.) Instantaneous One-zero Also called ‘point sampling’ or sampling sampling ‘fixed-interval time point sampling’ Score – proportion of all sample points on which behaviour happened 32 (Martin and Bateson, 1986, Altmann, 1974) Time sampling/interval structured recording – intervals & sampling points Sampling intervals need to be determined in pilot studies. 33  Rules for data recording Recording rules (when) Continuous Time = Interval structured recording recording sampling At pre-determined intervals (every 30 sec, every 5 min etc.) = time point sampling Also called ‘point sampling’ or Instantaneous One-zero ‘fixed-interval time point sampling’ sampling sampling Suitable for behaviours that last longer (states) than the sampling interval Score – the proportion of all sample points on which behaviour happened 34 (Martin and Bateson, 1986, Altmann, 1974)  Rules for data recording Recording rules (when) Continuous Time recording sampling During pre-determined intervals (every 30 sec, every 5 min etc.) Did the behaviour occur – ‘yes = One’ or ‘no = Zero’ Instantaneous One-zero sampling sampling Also called ‘fixed-interval time span sampling’ number of intervals that includes the behaviour Score – the proportion of all sample points on which behaviour happened 35 (Martin and Bateson, 1986, Altmann, 1974) Rules for data recording 36 (Broom and Fraser 2008) Rules for sampling Sampling rules (who, what) Ad lib Focal Scan Behaviour sampling sampling sampling sampling Ad libitum – ‘freely, as much as you desire, without restriction’ States and events Often used in field studies when generating an ethogram 37 (Martin and Bateson, 1986, Altmann, 1974) Rules for sampling Sampling rules (who, what) Ad lib Focal Scan Behaviour sampling sampling sampling sampling Continuously following one specific individual. States and events Often used for time budgets and social interactions. A sample of focal animals provides behavioural insights into large groups https://www.youtube.com/watch?v=ay-7LNMab40 38 (Martin and Bateson, 1986, Altmann, 1974) Rules for sampling Sampling rules (who, what) Ad lib Focal Scan Behaviour sampling sampling sampling sampling Group of animals is scanned at the same time Usually states Often used for time budgets, synchrony 39 (Martin and Bateson, 1986, Altmann, 1974) Rules for sampling Sampling rules (who, what) Ad lib Focal Scan Behaviour sampling sampling sampling sampling Specific behaviours of interest are recorded Usually events Also called ‘all occurrence’ or ‘conspicuous behaviour’ sampling 40 (Martin and Bateson, 1986, Altmann, 1974) Methods can be (and are often!) combined Sampling rules (who, what) Ad lib Focal Scan Behaviour sampling sampling sampling sampling Recording rules (when) Continuous Time recording sampling Instantaneous One-zero sampling sampling 41 (Martin and Bateson, 1986, Altmann, 1974)  Continuous recording 0 Latency (e.g. 1 min before behaviour starts) 30 min Behaviour 3 2 10 Continuous Total number of occurrences = 3 Frequency = 3 / 30 = 0.1 times per minute Duration = 3 + 2 + 10 = 15 min Mean duration = 15 / 3 = 5 min Proportion of time = 15 / 30 min = 0.5 (or 50% of the time) 42  Time sampling 0 30 min Behaviour 3 2 10 Proportion time Continuous 15 / 30 = 0.5 0 5 10 15 20 25 30 min Instantaneous Score 2 / 6 = 0.33 x x 0 5 10 15 20 25 30 min One-zero Score x x x x x 5 / 6 = 0.83 Behaviour and sampling methods will influence your results! 43 Your turn How much time hens spent in different locations during a day? (litter, perch, nest box) Ad lib Focal sampling sampling Scan Behaviour Continuous Instantaneous One-zero sampling sampling recording sampling sampling 44 Your turn Number of times birds peck at each other during 10 min? Ad lib Focal sampling sampling Scan Behaviour Continuous Instantaneous One-zero sampling sampling recording sampling sampling 45 Training is crucial Did you notice anything else? 46 https://youtu.be/ay-7LNMab40?si=0UAgxyOIAIuYIiNP Student resource –15-minutes Animal Behaviour sampling video 47  ANSC*3090 – 17 October 2024 1 The scientific method (adapted from Malmfors, Gransworthy, Grossman, 2004) 2 How to develop a research question? https://learningcommons.lib.uoguelph.ca/item/developing- research-question-worksheet 3 How to develop a research question? Chopped straw on the ground Visiting a friend’s backyard chickens….. 4  How to develop a research question? “Descriptive” research question: On average, how much straw do hens use to build their nests in the nest boxes? https://www.trentu.ca/academicskills/how-guides/how-succeed-math-and-science/writing-lab-reports/understanding-hypotheses-and-predictions 5 How to develop a research question? “Causal” research question: Why do hens prefer to make their nests in nest boxes with chopped straw instead of grass? https://www.trentu.ca/academicskills/how-guides/how-succeed-math-and-science/writing-lab- reports/understanding-hypotheses-and-predictions 6 What is a hypothesis? A proposed explanation of a pattern or phenomenon that may or may not be true Tentative We test the hypothesis using specific observations or Testable experiments, to provide evidence for or against it Falsifiable There can be more than one hypothesis under test at once (‘competing’ hypotheses) 7 What is a hypothesis? ……, you might hypothesize, “Hens use straw in their nest boxes instead of grass because straw is more abundant on the ground in their chicken coop.” Chopped straw on the ground 8 What makes a “good” hypothesis? A verifiable, testable statement; Very clear and specific Postulates connections between cause and effect about the observed outcome A study can never prove hypotheses, as alternative explanations cannot be entirely ruled out. Hypotheses can only be rejected = falsified, or deemed to remain valid. How? Makes specific predictions that can be tested with data, and that distinguish it from other competing hypotheses 9 What makes a “good”, “explanatory “ hypothesis that explains the observed behaviour? Hypothesis H1 (working/alternative hypothesis) Associated with the research question, predicts that there is a difference between samples Hypothesis H0 (null hypothesis) Predicts there is no significant difference between the samples. In research/practice: H0 is not explicitly stated 10 Example: Chickens in my neighbour’s garden…. 11  What makes a “falsifiable” hypothesis? Hypothesis H1 (working/alternative hypothesis) There are chickens in my neighbour’s garden. The alternative hypothesis (H1) can be verified but not falsified, as you might not have checked thoroughly enough or at the correct time to find chickens there. Hypothesis H0 (null hypothesis) There are no chickens in my neighbour’s garden Once chickens are observed in my neighbour’s garden, the hypothesis is falsified. 12 What makes “good” hypotheses and predictions? A hypothesis leads to predictions Predictions: outcome you observe if your hypothesis is correct Predictions: Determine which behaviours/physiological measurements are collected; “if-then” statements 13  Predictions, following the nest material example…. If hens use chopped straw because it is more abundant, and we compare areas with more grass than chopped straw, then nests should be made out of grass. … REFINED: If hens choose nesting materials based on their abundance, then when grass is more abundant, hens will use those in their nests 14 Hypothesis vs Prediction Hypothesis ≠ Prediction (but predictions follow from hypothesis) Why are nests of hens made with chopped straw rather than grass? Hypothesis: My neighbour’s hens use chopped straw in their nests rather than grass because chopped straw is the more abundant material in their environment (abundance hypothesis) Prediction: “If my neighbour’s hens chooses nesting materials based on their abundance, then when grass is more abundant, hens will use grass in their nests.” 15 Tinbergen noticed that the inside of the egg is not camouflaged... 16 Hypothesis: “Remove eggshells from their nests to reduce predation risk” If this idea is correct, what should we see? Predictions: If Black-headed Gulls remove eggshells promptly after hatching, then fewer predators will be attracted to the nest area, increasing the survival rate of their chicks. 17 Hypothesis: “Remove eggshells from their nests to reduce predation risk” 18 22 October 2024 1 Observational Studies - Field work https://youtu.be/-7caGCKECO4?si=THN7rDyDRDTu4xOw 2  Observational Studies – Field work Domesticated – wild animals Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize in Physiology or Medicine with Nikolaas Tinbergen and Karl von Frisch. Pioneers of the study of Dr. David Wood-Gush (1922-1991) ethology, inspiring generations 3  What are domestic pigs? Descended from the wild boar Omniverous forest dwellers Spend most of their active time foraging 4  How do domesticated pigs behave in the habitat of their wild ancestors? Studied the “natural” behaviour of domestic pigs in a semi-natural environment, the Edinburgh “Pig Park”. 5 How do domesticated pigs behave in the habitat of their wild ancestors: Wood-Gush & Stolba Time budgets Time-specific observations e.g. circadian rhythm Situation-specific observation e.g. predator Behavioural development They studied “natural behaviour” in a “semi-natural environment” in the 1980ies! 6 Experience of the observer Are you measuring the same thing over time? 7 Observer Biases Experimenter Bias 8  Observer Biases Expectation Bias Rosenthal effect/ experimenter expectancy effect Experiment: Students were told they were working with either “maze-bright & maze-dull” rats ALL rats were the same = student’s expectations affected the outcomes 9  Observer Biases Expectation/Confirmation Bias The behaviour of the animal is not influenced directly favour data that supports hypotheses remember information that confirms existing beliefs Record behaviour in such a way that supports the expectations/beliefs 10 Recent example: Observer bias Humans cannot be assumed always to process information objectively and accurately 11 Observer bias ~160 3rd year veterinary students Shown different video clips (pigs, cows, chickens) Students were told different background information about the animals 12 (Tuyttens et al., 2014) Observer bias Different results for behavioural observations SBV+ group was told that pigs were selected for high social breeding value 13 Observer bias and “blinding” Ways around this? Making sure the observer is unaware of treatments Making sure the person analyzing the data is unaware of the treatments Minimize the influence of observers to make subjective decisions = detailed and mutually exclusive ethogram Experienced versus naïve observer Multiple observers – observer effects – agreement between observers 14 Observer reliability Intra-observer reliability – within an observer Are you measuring the same thing over time? Not always reported in papers! Inter-observer reliability – between observers Are you measuring the same thing as the other observer(s)? It is crucial that observers consistently score/categorize the same states/events in a similar manner. 15 Two observers in an observational study Observation Example: Two observers, Observer A Period Observer A Observer B and Observer B, are recording 1 Yes Yes instances of social interactions over 10 2 No No observation periods. 3 Yes Yes 4 No Yes Each observer notes whether the 5 Yes No behaviour occurred (Yes) or did not 6 No No occur (No) during each period. 7 Yes Yes 8 No No 9 Yes Yes e.g. Kappa (Cohen, 1960) index for 10 No No measuring inter-observer agreement 16 Calculating Cohen’s Kappa – step 1 Contingency Table Observer B Observer B Total Yes No Observer A 5 1 6 Yes Observer A 1 4 5 No Total 6 5 11 Calculate the Observed Agreement (Po): Po = (Number of agreements) / (Total observations) Po = (5 + 4) / 10 = 0.9 17 Calculating Cohen’s Kappa – step 2 Contingency Table Observer B Observer B Total Yes No Observer A 5 1 6 Yes Observer A 1 4 5 No Total 6 5 11 Calculate the Expected Agreement (Pe): Pe = [(Row total for Yes * Column total for Yes) + (Row total for No * Column total for No)] / (Total observations)2 Pe = [(6 * 6) + (5 * 5)] / 10^2 = (36 + 25) / 100 = 0.61 18 Calculating Cohen’s Kappa – step 3 Contingency Table Observer B Observer B Po = (5 + 4) / 10 = 0.9 Total Yes No Observer A 5 1 6 Yes Pe = [(6 * 6) + (5 * 5)] / 102 = (36 + 25) / 100 = 0.61 Observer A 1 4 5 No Total 6 5 11 Calculate Cohen’s Kappa (κ): κ = (Po - Pe) / (1 - Pe) κ = (0.9 - 0.61) / (1 - 0.61) = 0.29 / 0.39 ≈ 0.74 19  Observer Biases Clever Hans Effect/Phenomenon Unconscious movements/unintentional cues Influence the behaviour of the animal 20 Sampling Biases Selection of study animals: Which focus animal represents the entire population? 21 Sampling Biases STRANGE framework for identifying “Sampling bias” Webster & Rutz 2020 Social background Trappability and self-selection Rearing history Acclimation and habituation Natural changes in responsiveness Genetic make-up Experience 22 Sampling Biases Social background Trappability and self-selection Rearing history 23 Sampling Biases Acclimation and habituation Natural changes in responsiveness Genetic make-up Experience 24 Resources needed? Technology? Live recording – paper, pen, clipboards Video/audio recording – cameras, audio recorders, storage 25 Video and audio recording devices: Data in greater detail Legend A) Jump tower B) Jump tower platform C) Lateral camera D) Frontal camera 26 Example – High Speed Video Camera 27 Frontal Video Wing Beat Frequency (Hz) Wing Beat Amplitude (Rad) Angular Velocity (Rad/s) Wing Beat Frequency Asymmetry Wing Beat Amplitude Asymmetry Angular Velocity Asymmetry 28 High-speed video camera – lateral view 29 Lateral Video Descent Angle (°) Acceleration (m/s2) Legend A) Beak point 1 Resultant Velocity 1 (m/s) B) Beak point 2 C) Beak point 3 Resultant Velocity 2 (m/s) D) Jump tower E) Jump tower platform Descent Velocity (m/s) F) Meter stick 30 Example: Video analysis tool – Mangold INTERACT Overview 31 Resources needed? Technology? Validation! 32 Examples Data loggers for animal behaviour and animal physiology: Locomotion Transmitter-Receiver Systems Radio-Frequency Identification (RFID) - passive transponders Global Positioning System (GPS) – active transmitters Animal Tracking 33 People vs Technology Observer Validation! reliability! (Ellen et al., 2019) 34  RFID-equipment Equipment Leg Bands (Garant et al., 2022) 35 Antenna Placement Animal Tracking – activity changes at pixel-level (example) EthoVision XT: The video tracking software 36 Automated data recording and analysis tools (example) TrackLab- System for intelligent tracking analysis https://youtu.be/fru-j5OmwFo?si=ewUXBoAsLNTMiNcL 37 Activity levels – accelerometers Understanding output data & validation Data- logger 38  Activity Levels – Accelerometers Low-intensity physical activity No change in location, postural movements such as sleep-like-resting, neck-shortening-resting, sitting, sleeping, minor head/shoulder/neck movements Moderate-intensity physical activity Small change in location, foraging, walking, preening, drinking, brief wing movements, Accelerometer with 16-bit ADC. Running at 10-100 Hz and 8g’s, recording acceleration in X, Y and Z direction and measuring atmospheric pressure. searching High-intensity physical activity Usually a change in location, walking, running, controlled aerial ascent/descent, shaking, jumping 98% accuracy verified using 39 video Example: Force Plates Ground reaction forces (GRF) of the foot increased gradually with increasing slope angle Higher GRFs and longer ground contact times force- plate were recorded for the first preparatory step for steepest inclines Domestic fowl modulate locomotion patterns in response to incline angle in a manner that suggests anticipation Data on kinetic modulation are important to understand how birds locomote their environment in a controlled and safe manner Installed cameras around the test arena to observe and record behaviour without needing to be physically present OUTSIDE TEST ARENA 41 INSIDE TEST ARENA – remote monitoring 42 Training to jump on force plates from 30 cm platform to obtain a treat A digital revolution (examples) Radio-frequency identification (RFID) Accelerometers Force plates Rumination / activity collars (Moreau et al., 2009) Cough monitors Technology always needs to be validated (Ellen et al., 2019) 44  A digital revolution Precision livestock farming use of sensors/programs to manage livestock Automatic milking systems for cows Electronic feeding stations for sows Automatic weighing stations for chickens https://www.youtube.com/watch?v=62ioHr8iTu8&t=1s https://dairy-360.uoguelph.ca/- for more 45 (Stygar et al., 2021; Tuyttens et al., 2022)  A digital revolution Keep in mind most technologies not designed with purpose to improve animal welfare 46 (Stygar et al., 2021; Tuyttens et al., 2022) 24 October 2024 1  What we discussed so far….. # Non-Experimental Observations * Time budgets Nauralistic observations observing in the wild - Behavioural Development over time looking - at normal behaviour Time and situation-specific observations Goal: understanding the behaviour in its natural context in real- world settings 2 Cameras, sensors, software needed for non-experimental and experimental observational data recording Also require ethical approval protocols ? - , - Transportation ? -Funding planning + 3  Equipment needed? Technology?... cameras, sensors , software 1. Live recording – paper, pen, clipboards 2. Video/audio recording – cameras, audio recorders, storage 4 Video audio recording “Picking grapes from the vine” 5 Video analysis - picking grapes I. Manual analysis: pen, paper, Excel spreadsheet or similar software 2. Video Analysis Software: collecting, analyzing and presenting observational data 6 Steps for Video Analysis in Behavioural Studies Video Import: Import video files into the software. Coding Scheme Setup: Define the states or events you will be looking for in the video. Annotations: Manual Annotations: Use the mouse or keyboard to log behaviors. For example, specific keys can be assigned to different behaviors (e.g., pressing Key F logs “Foraging” along with the timestamp). Automated Annotations: The software automatically detects and logs behaviors. Data Analysis & Visualization: Analyze data for duration, frequency, synchronization, and generate figures and tables. 7  Video analysis tools – commercial and open-source software Example: Mangold INTERACT Overview 8 Examples of tracking locomotion using videos, sensors and software 9 Video Piglets outdoor locomotion 10  Automated data recording and analysis tools: e.g. Ultra-Wideband (UWB) technology Noldus: TrackLab- System for intelligent tracking analysis https://youtu.be/fru-j5OmwFo?si=ewUXBoAsLNTMiNcL 11 Animal Tracking – activity changes at pixel-level (example) EthoVision XT: The video tracking software 12 Video and audio recording devices: Data in greater detail Legend A) Jump tower B) Jump tower platform C) Lateral camera D) Frontal camera 13 Example Test arena: High-Speed Video Camera to capture wing movements 14 Frontal Video Wing Beat Frequency (Hz) Wing Beat Amplitude (Rad) Angular Velocity (Rad/s) Wing Beat Frequency Asymmetry Wing Beat Amplitude Asymmetry Angular Velocity Asymmetry Frame-by-frame analysis offers a detailed view of wing movements, aiding in the understanding of flight biomechanics. 15 Test arena: High-speed video camera – lateral view 16 Lateral Video Descent Angle (°) Acceleration (m/s2) Legend A) Beak point 1 Resultant Velocity 1 (m/s) B) Beak point 2 C) Beak point 3 Resultant Velocity 2 (m/s) D) Jump tower E) Jump tower platform Descent Velocity (m/s) F) Meter stick 17 Resources needed? Technology? Validation! 18 Mounting data loggers on animals - providing data on animal behaviour and movement Transmitter-Receiver Systems Radio-Frequency Identification (RFID) - passive transponders Global Positioning System (GPS) – active transmitters Animal Tracking 19 People vs Technology Observer Validation! reliability! (Ellen et al., 2019) 20 RFID-equipment – two-dimensional tracking Equipment Leg Bands (Garant et al., 2022) 21 Antenna Placement Activity levels – Triaxial accelerometers Understanding output data & validation Data- logger Home pen 22  Activity Levels – Accelerometers Low-intensity physical activity No change in location, postural movements such as sleep-like-resting, neck-shortening-resting, sitting, sleeping, minor head/shoulder/neck movements Moderate-intensity physical activity Small change in location, foraging, walking, preening, drinking, brief wing movements, Accelerometer with 16-bit ADC. Running at 10-100 Hz and 8g’s, recording acceleration in X, Y and Z direction and measuring atmospheric pressure. searching High-intensity physical activity Usually a change in location, walking, running, controlled aerial ascent/descent, shaking, jumping 98% accuracy verified using 23 video Automated data recording and analysis tools: e.g. Two-axial accelerometer- based pedometers in research & practical applications 24  Example: Force Plates to understand and improve walking patterns Ground reaction forces (GRF) of the foot increased gradually with increasing slope angle force- Higher GRFs and longer ground contact times plate were recorded for the first preparatory step for steepest inclines Domestic fowl modulate locomotion patterns in response to incline angle in a manner that suggests anticipation Data on kinetic modulation are important to understand how birds locomote their environment in a controlled and safe manner Installed cameras around the test arena to observe and record behaviour without needing to be physically present OUTSIDE TEST ARENA 26 INSIDE TEST ARENA – remote monitoring 27 Training to jump on force plates from a 30 cm platform to obtain a treat gait and balance testing in animals with musculoskeletal issues A digital revolution (examples) Radio-frequency identification (RFID) GPS tracking collar HOBO® Pendant G tri-axial accelerometer Accelerometers Force plates Rumination / activity collars (Moreau et al., 2009) Cough monitors Technology always needs to be validated Ultra-wideband vs video tracking (Ellen et al., 2019) 29  Positioning measuring devices on animals How do you choose the correct position? What are your research goals? E.g. overall vs. head movements Potential for device movement vs. too tight? Weight of the device? Does the device interfere with the animal’s species-specific behaviour? Is it accurate for detailed locomotion analysis? 30  A digital revolution Precision livestock farming use of sensors/programs to manage livestock Automatic milking systems for cows Electronic feeding stations for sows Automatic weighing stations for chickens https://www.youtube.com/watch?v=62ioHr8iTu8&t=1s https://dairy-360.uoguelph.ca/- for more 31 (Stygar et al., 2021; Tuyttens et al., 2022)  A digital revolution Keep in mind most technologies not designed with purpose to improve animal welfare Direct harm to animals; indirect effects via end-user; changes to housing and management; ethical stagnation/degradation (Stygar et al., 2021; Tuyttens et al., 2022) 32 Experimental observations Observing animals in their home versus test environments 33 Why do we observe behaviour in artificial test environments? 34 Behavioural tests Behavioural tests can be used to answer specific questions in an optimal way Often a proxy, often faster and more accurate than time- consuming direct behavioural observation Can be applied to different species (often developed from rodent studies) and measure different aspects of the animal (Nielsen, 2020) 35  Common tests for animal behaviour (examples) Research area Behavioural test Activity and exploration Locomotor activity/rearing, open field test, hole-board test, response to novelty, voluntary exercise Anxiety Open field test, locomotor activity/rearing, elevated plus maze tests, dark/light test Depression Forced swim test*, tail suspension test Learning and memory Passive or active avoidance, Aron test, Morris water maze, radial maze test, T-maze test, object recognition test, operant procedures, 5/9-hole test Reward Place preference tests, operant procedures Sensory and motor skills Coordination and equilibrium, grip strength, exercise training, nociception, startle response (Nielsen, 2020) Social behaviour Social interaction, social reinstatement 36 Specialized equipment used for behavioural tests (examples) 37 What can you do with behavioural tests? Tests to characterize the animal Choice, preference and motivation Ability to detect and distinguish Effects of age and treatment Reinforcement and punishment Learning capacity, memory, and cognitive ability Genetic components of behaviour (Nielsen, 2020) 38 Individual differences characterize an animal Genetic, development & current environment 39 Individual differences Example video: Calves exhibit a range of behavioural patterns when encountering a new object - Individual differences - Past experiences - Current circumstances Exposing individuals to personality tests 40 Tests to characterize animals 41  Open Field Test Initially developed and validated for rats (nocturnal animals) “Fear-testing” Outcome variables in the bright open space: Total distances moved/time spent moving Frequency of rearing Time spent in the center vs. periphery Defecation rates Periods of immobility Frequency and duration of grooming, sniffing Duration for an open-field test: 5-10 min 42 Open- Field Test (Nielsen, 2020) Quail in an Open Field Test 43  Ancestors of cattle lived in open grasslands and Open Field Test - forested areas….. When testing other species….pigs, cattle, horses, etc… Arena dimensions Timing Outcome variables Inclusion of additional tests – novel object Inconsistent set-up 44  Tonic Immobility (Nielsen, 2020) Tonic immobility (TI): unlearned response induced by physical restraint; catatonic-like state; reversible, reduced responsiveness to external stimulation; evolutionary adaptive behaviour; defence reaction against predator attack. 45 29 October 2024 1 What can you do with behavioural tests? Tests to characterize the animal Choice, preference and motivation Ability to detect and distinguish Effects of age and treatment Reinforcement and punishment Learning capacity, memory, and cognitive ability Genetic components of behaviour (Nielsen, 2020) 2 Individual differences characterize an animal Genetic, development & current environment 3 Individual differences Example video: Calves exhibit a range of behavioural patterns when encountering a new object - Individual differences - Past experiences - Current circumstances Exposing individuals to personality tests 4 Tests to characterize animals Interestingly, there is a focus on fear Why? 5  Panksepp’s “Basic Emotions” https://www.amazon.com/Expression-Emotions-Man-Animals/dp/1470188880 6 Darwin, 1872 Panksepp, 1998 The Expression of the Emotions in Man and Animals Affective Neuroscience. The Foundation of Human and Animal Emotions Historically, emotion research focused on negative emotions…. Consistent behavioural reactions Clear physiological markers Evolutionary significance This can be induced in the laboratory setting 7 Figure 11.1 Components of an Emotional Response Role of amygdala & prefrontal cortex Figure 11.2 Amygdala Projections Three major regions of the amygdala: Lateral Nucleus: Receives information from the neocortex, thalamus, and hippocampal formation. Basal Nucleus: Receives information from the lateral nucleus, ventral striatum, and thalamus. Central Nucleus: Receives information from the lateral and basal nuclei and projects to the hypothalamus, midbrain, pons, and medulla. Activation of the central nucleus leads to behavioural, autonomic, and hormonal responses. 9  Behavioural and physiological responses of the central nucleus of the amygdala Figure 11.3 The Outputs of the Central Nucleus of the Amygdala Sympathetic Nervous System Activation: Increased heart rate; Increased blood pressure; Paleness Parasympathetic Nervous System Activation: Urination; Defecation; Increased respiration Behavioural arousal (increased alertness, readiness to act, attentive) Increased vigilance Cortical activation Startle response (exaggerated reaction to unexpected stimulus (loud noise/sudden movement) Behavioural arrest (freezing) Facial expression of fear ACTH and glucocorticoid secretion Cortical activation (crucial for attention, memory, emotional 10 responses)  Open Field Test Initially developed and validated for rats (nocturnal animals) “Fear-testing” Outcome variables in the bright open space: Total distances moved/time spent moving Frequency of rearing Time spent in the center vs. periphery Defecation rates Periods of immobility Frequency and duration of grooming, sniffing Duration for an open-field test: 5-10 min 11 Open- Field Test (Nielsen, 2020) Quail in an Open Field Test 12  Ancestors of cattle lived in open grasslands and Open Field Test - forested areas….. When testing other species….pigs, cattle, horses, etc… Arena dimensions Timing Outcome variables Inclusion of additional tests – novel object Inconsistent set-up 13  Open Field Test Arena Novel Object Recognition and Object Location Behavioral Testing in Mice on a Budget Denninger, J. K., Smith, B. M., Kirby, E. D. Novel Object Recognition and Object Location Behavioral Testing in Mice on a Budget. J. Vis. Exp. (141), e58593, doi:10.3791/58593 (2018). 14 Open Field Test Arena Building Video https://app.jove.com/v/58593/novel-object-recognition-and- object-location-behavioral-testing-in-mice-on-a-budget 15 Fear induced behaviour Fear of humans 17 Giraffe, leopard, hyena, zebra, kudu, warthog and impala running in response to hearing humans Giraffe, leopard, hyena, zebra, kudu, warthog and impala running in response to hearing humans 18 Defensive Responses & Threat 19 Immobility & Predation Forceplate 20  Tonic Immobility (Nielsen, 2020) Tonic immobility (TI): unlearned response induced by physical restraint; catatonic-like state; reversible, reduced responsiveness to external stimulation; evolutionary adaptive behaviour; defence reaction against predator attack. 21 Tonic Immobility Test The test uses a V-shaped crate or a table (e.g., for birds). The animal is turned on its back, and a lightweight (such as a hand or bag) is placed on its chest. Outcomes: - Number of inductions - Latency to enter tonic immobility (TI) - Duration of TI - attempt to right itself “Backtest” in pigs 22  Novel – Object Test Leverage rodent’s innate curiosity to explore New stimuli without reinforcement or punishment Performed after Open Field Test or in the same arena as the open field Individual/groups; arena/home pen Objects: Bright, novel shape Creates motivational conflict in the animal; making it both fearful and curious about the object 23 Novel – Object Test Outcomes: latency to the first contact; number of object contacts; handling intensity (e.g. tasting, biting, eating, etc); fear-related behaviour in the arena 24 Startling stimulus Novelty, unpredictability, and suddenness help predators increase their chances of a successful hunt. Observing an animals reaction to Reaction to novelty: e.g. presenting a closed orange umbrella Reaction to suddenness: e.g. Opening of an orange umbrella e.g. Erhard et al. 2004 (sheep); 25  Startle Test Startle test: exposure to a sudden and intense sound or visual stimulus The startle response is enhanced when a conditioned fear stimulus is presented before the actual stimulus Conditioned fear test: exposure to a signal that was previously associated with nociceptive (pain) or disturbing event (electric shock) 26 Startle Test Nikon Speedlight SB-20 xenon camera flash Forceplate JVC GG-PX100 600 frames/sec © MISHA ROSS © MISHA ROSS 27 Human Approach Test Forced approach test – the animal is approached by a human - more likely to elicit an active response Voluntary approach test – the animal is free to approach the experimenter - more likely to elicit passive response Silhouettes of predators “Hawk-goose phenomenon” 29  Resident-Intruder Model (rat example) Resident male rats will attack unfamiliar males that intrude in its home cage To determine the violent nature – different types of intruders (females, anesthetized males) or a novel environment is used To determine context – a detailed quantitative analysis of the offensive behavioral repertoire is required Real world scenario: Based on aggressive scores from resident-intruder tests, attack latency can be used to predict how pigs will behave in a group when a stressor, such as mixing familiar with unfamiliar pigs, occurs (D`Eath, 2002) Resident-intruder Paradigm http://www.jove.com/video/4367/the-resident-intruder- paradigm-standardized-test-for-aggression Defensive Behaviours and Predation Figure 11.7 Neural Circuitry in Defensive Behavior and Predation Aggressive behaviour: threatening gestures and/or attacks Species-specific movements such as posturing, biting, hissing, etc. Threatened animals can show: A. defensive behaviours B. submissive behaviours Threatening behaviours help reinforce social hierarchies, warning intruders away from an animal territory; not involving fighting = avoid harm; increased survival Attacking a member of the same species: activity of sympathetic NS is high Attacking a member of a different species = PREDATION The activity of the sympathetic NS is low; predator is not angry with prey 32  Signal input: Signal input: Hunger, thirst, Pain, sudden noise, frustration sudden movement lateral medial Hypothalamus Hypothalamus Glutamat midbrain brainstem Siegel & Victoroff 2009  The limbic systems receives critical inputs from the cortex, sensory signals, which provides such signals with emotional qualities, which then modulate functions of the hypothalamus and midbrain (PAG).. This takes place because when Reciprocal one of the responses is prepotent, inhibitory the other is suppressed connections by virtue of the between LH and MH inhibitory relationship with utilize GABA as a the other region. neurotransmitter. The relationship allows either predatory attack or defensive rage to Little Approach Sympathetic occur at a given activation locomotion time. Target specific Sympathetic activation flight locomotion Not target specific Siegel & Victoroff Siegel & Victoroff 2009 2009 Many more tests….. - Elevated Plus Maze - Cognitive Bias - Etc. 35 Apply your knowledge: A series of test procedures can yield similar or varying behavioural and physiological results Model animal: Japanese Quail 36  Requires minimal space Cost-effective Short generation interval (3-4 months) Excellent model for studying chickens Japanese quail Coturnix japonica  TI: genetic Tonic Immobility (seconds) 300 250 LTI 200 150 100 Control (unselected) 50 STI 0 10 20 30 Generations Figure: 40 generations of selection for Long (LTI) and Short (STI) Tonic Immobility in Japanese Quails  TI: age and sex The duration of tonic immobility was consistent across both sexes and was not influenced by the age of the birds. 39 Open Field Test: enrichment & handling STI chicks showed less freezing, vocalized earlier, and walked more frequently and sooner in an open field. Environmental enrichment decreased silence and inactivity, while regular handling increased vocalization and shortened TI duration. 40 Hole-in-the-Wall Test of Timidity / Emerge Test wooden box with a hinged lid & guillotine trapdoor covering an opening in one end wall placed in a brightly lit open field. STI chicks emerged sooner from the box, and environmental enrichment accelerated their emergence. 41 Genetic selection for less fearful quails made them easier to catch 42 HPA-axis responsiveness: Fear & stress responses are closely linked - no line differences in basal plasma corticosterone (CORT) levels - restraint in a crush cage: 10 minutes – caused sign. increase in CORT in both lines. Crush cage - the increase was sign. greater in STI quails. This difference persisted for at least 2 hours (Hazard et al. 2005). 43 General Behavioural Test Considerations Manual testing and handling: prior handling, the experience of the handler, etc. Testing environment: isolation of animal; separate room, arena, etc., lighting, sound, odour etc.; time of the day, etc. Protocol variability: equipment used; definition of behavioural outcomes, habituation, etc.; multiple tests in a row; etc. Animal: strain (genetic); sex, age, physiological states, disease/health, home environment, prior experience of the animal, etc. 44 31 October 2024 1 What can you do with behavioural tests? Tests to characterize the animal Learning capacity, memory, and cognitive ability Reinforcement and punishment Choice, preference and motivation Ability to detect and distinguish Effects of age and treatment Genetic components of behaviour (Nielsen, 2020) 2 3 Importance Aggression, violence and social stress are serious problems in our human society & in animals kept in human managed systems Hence, there is a need to understand these behaviours in terms of their underlying neural/hormonal causal mechanisms  body lesions in the loose herds vulva lesions caused by biting Aggression between sows after mixing Inter-female aggression in breeding rabbits Skin injuries –I Group housing of does with pups indicator for aggression among does  Aggression is normal It can be highly functional – as social communication aimed at active control of the social environment: Establishment of a territory Social dominance Defence of resources Aggression is most likely…. When the interests of two or more individuals are in conflict E.g.: can be sensitive to spacing between individuals Benefit: get what you want Costs: consumes a time & energy budget reduces vigilance for predators risk of injury (ultimate cost=death) Aggression and physical conflicts Are potentially harmful for both parties involved Therefore, adaptive control mechanisms develop to minimize and control physical aggression Such mechanisms include: Threatening behavior Taboos Ritualization Submission Reconciliation Appeasement  Types of aggression Categories of evolutionary purpose (Moyer 1968) 10  Unrelated to the other forms of aggression cat Attack response is triggered by the presence of a prey object within its visual field mouse Predatory aggression Fear induced aggression Aggression induced when an animal is prevented from escape and attacks another animal E.g. Dog aggression is a serious problem. It appears that fear- induced aggression in E-Shock collars induce fear, dogs is often discomfort and pain! implicated.  Maternal - aggression Aggression of a mother when it perceives that its offspring is threatened e.g. Broody hen defends eggs and chicks as best, she can. She makes her feathers big and fluffy Inter-male aggression (see Resident-Intruder video) Induced when a male is paired with a male of the same species It could be due to a variety of reasons: Territory Mating Dominance, etc. Territorial aggression Donkeys use front and hind legs as well as their teeth to bite intruders. Induced following the entry of an animal (intruder) into the domain deemed to be established by another animal (resident) = resident-intruder model  ©APA/EPA/TOL GA BOZOGLU Territorial aggression/Territorial wars Classification of aggression - Dichotomous classification schemes Offensive/proactive/predatory aggression Defensive/reactive/affective/impulsive aggression/defensive rage Aggression turning into violence (see Resident-Intruder-video) Violence: an injurious form of offensive aggression that is out of control and out of context It is a pathological form of offensive behavior – no longer subject to inhibitory control mechanisms It has no additive functional value in social communication (de Boer& Koolhaas 2005)  Violence Differs both quantitatively and qualitatively from normal adaptive Growing rabbits/ offensiveness scrotum injuries May include bites, attacks at vulnerable body parts (e.g. throat, belly, paws)  Resident-Intruder Model (rat example) Resident male rats will attack unfamiliar males that intrude in its home cage To determine the violent nature – different types of intruders (females, anesthetized males) or a novel environment is used To determine context – a detailed quantitative analysis of the offensive behavioral repertoire is required Real world scenario: Based on aggressive scores from resident-intruder tests, attack latency can be used to predict how pigs will behave in a group when a stressor, such as mixing familiar with unfamiliar pigs, occurs (D`Eath, 2002) Resident-intruder Paradigm http://www.jove.com/video/4367/the-resident-intruder- paradigm-standardized-test-for-aggression Physiology of aggression Aggression is based on brain mechanisms Learning Hormones Defensive Behaviours and Predation Aggressive behaviour: threatening gestures and/or attacks Figure 11.7 Neural Circuitry in Defensive Behavior and Predation Species-specific movements such as posturing, biting, hissing, etc. Threatening behaviours help reinforce social hierarchies, warning intruders away from an animal territory; not involving fighting = avoid harm; increased survival Threatened animals can show: PAG= A. defensive behaviours Periaqueductal gray / MIDBRAIN B. submissive behaviours Attacking a member of the same species: activity of sympathetic NS is high Possible connection between lateral hypothalamus and ventral PAG not verified. Attacking a member of a different species = PREDATION The activity of the sympathetic NS is low; predator is not angry with 23 prey 24 The limbic system receives critical inputs from the cortex and sensory signals, which provide signals with emotional qualities and modulates functions of the hypothalamus and midbrain (PAG).. Reciprocal This takes place inhibitory because when connections one of the between LH and MH responses is utilize GABA as a prepotent, neurotransmitter. the other is The relationship suppressed allows either by virtue of the predatory attack or inhibitory defensive rage to relationship with occur at a given the other region. time. Approach Little locomotion Sympathetic activation Target specific Sympathetic flight locomotion activation Siegel & Victoroff 2009 Not target specific Siegel & Victoroff 2009 Predatory aggression/attack behavior The response is highly directed to the prey/target Little sympathetic activation This response requires planning and a strategy (Siegel 2005) Positive reinforcement  Defensive behaviour Increases in sympathetic activation (fight-or- flight) Response is impulsive & lacks cortical involvement Displacement of the target to other individuals in the environment (in contrast to predatory attack- target is highly specific) Motivational mechanisms set of homogeneous neurons Motivational mechanisms  set of homogeneous neurons Hormonal action Offensive animal (if wounded at all) is bitten on the face Motivational Learning takes place mechanisms Defensive animal is wounded on flank and rump Many more tests to characterize animals….. - Elevated Plus Maze - Cognitive Bias - Etc. 31 Apply your knowledge: A series of test procedures can yield similar or varying behavioural and physiological results Model animal: Japanese Quail 32  Requires minimal space Cost-effective Short generation interval (3-4 months) Excellent model for studying chickens Japanese quail Coturnix japonica  TI: genetic Tonic Immobility (seconds) 300 250 LTI 200 150 100 Control (unselected) 50 STI 0 10 20 30 Generations Figure: 40 generations of selection for Long (LTI) and Short (STI) Tonic Immobility in Japanese Quails  TI: age and sex The duration of tonic immobility was consistent across both sexes and was not influenced by the age of the birds. 35 Open Field Test: enrichment & handling STI chicks showed less freezing, vocalized earlier, and walked more frequently and sooner in an open field. Environmental enrichment decreased silence and inactivity, while regular handling increased vocalization and shortened TI duration. 36 Hole-in-the-Wall Test of Timidity / Emerge Test wooden box with a hinged lid & guillotine trapdoor covering an opening in one end wall placed in a brightly lit open field. STI chicks emerged sooner from the box, and environmental enrichment accelerated their emergence. 37 Genetic selection for less fearful quails made them easier to catch