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1

behaviour can be observed and measured to directly or indirectly identify underlying mechanisms - MAKE SURE TO LOOK AT NOTES WHILST DOING THESE

Causal or correlational evidence

2

Cambrian 'explosion' (542-520 MYA)

- Rise in diversity of organisms
- Rapid appearance of many animal body plans seen today
- Origin of vertebrates
- Development of the eyes
- Compound eyes that insects have - more complex than previously known
- Important in locomotion - leads to changes in the senses - capture info faster - outperform the prey
- 25mil years later = lens eye that we have

3

Cambrian explosion research

Runnegar (1981)

Wood et al. (2019)

4

Runnegar (1981)

- Appearance of fossils at end of Precambrian explained in 2 ways:
1. initial explosion of animal life
2. explosive appearance of fossilisable animals - supported by ev from oxygen-binding proteins of living animals - billion year history for lower invertebrate phyla
- Cambrian explosion caused by rise of large, muscular +/ mineralised animals - appearance explained by sharp increase in oxygen content of atmos

5

Wood et al. (2019)

- Cambrian explosion = rapid increase in animal diversity and abundance
- Nested within far more ancient record of macrofossils extending to late Ediacaran (571MYA)
- Evolutionary events documented within Ediacaran-Cambrian interval coincide with geochem ev for modernisation of Earth's biogeochem cycles
- Holistic integration of fossil and geochem records —> challenge that Ediacaran and Cambrian worlds distinct and places biotic and env change within longer term narrative
- Evolution of metazoans (an animal of the Metazoa division) may be fac by series of dynamic and global changes in redox conditions and nutrient supply - + biotic feedbacks = turnover events that sustained multiple phases of radiation
- Early metazoan diversification recast as series of successive, transitional radiations that extended from late Ediacaran —> palaeozoic
- Cambrian explosion just one phase in several metazoan radiations

6

mechanisms and adaptive functions of behaviour

- Proximate causation (during an animal's life time)
- Explanation of an animal's behaviour based on trigger stimuli and internal mechanisms (e.g. brain signals)
- How is behaviour produced?
- How does a behaviour develop?
- How is a behaviour inherited?
- Ultimate causation (over several generations)
- Explanation of an animal's behaviour based on evolution - why this specific trait was favoured by natural selection
- Why has a behaviour developed?
- What selective advantage does it provide?
- What is the evolutionary history?

7

mechanisms and adaptive functions of behaviour research

Otsuka (2015)

8

Ostsuak (2015)

- R'ship between proximate and ultimate causation ambiguous
- Causal graph theory - unified framework to systematically translate given 'proximate' causal structure into 'ultimate' evolutionary response
- Illustrates evolutionary implications of various causal mechanisms inc. epigenetic inheritance, maternal effects and niche construction
- Interplay between proximate and ultimate causation
- Formal method to eval/discover non-standard/yet unknown evolutionary phenom

9

Why care about (neuro)physiological mechanisms when studying behaviour

- Tinbergen's 4 explanatory levels as complementary ways of looking at behaviour (Tinbergen (1963))
- **proximate** - a description of the mechanisms underlying behaviour in an individual or group of animals
- **ontogeny** - development of behaviours within an individual lifespan
- **phylogeny** - development of behaviours over generations and in different populations
- **ultimate or functional** - the adaptvie significance or utility of the behaviour for the organism

10

Why care about (neuro)physiological mechanisms when studying behaviour research

Duckworth (2009)

Fink (1982)

11

Duckworth (2009)

- 2 difficulties in incorp behav as causal force in evolution
1. Focus on high plasticity of behav diverted attention from investigation of devel mechanisms - crucial for detecting novel behav patterns
2. Lack of clear evolutionary mechanisms - hindered understanding of how changes in behav that occur within indvs affect popn level processes
- Incorp devel and evolutionary mechanisms resolves issues and moves beyond Q of whether behav primarily drives/inhibits evolution - what circumstances behavs affect evolutionary processes?
- Recognition that behav can affect evolutionary change evident in models which are basis for popn genetics (Hardy-Weinberg) - random mate choice and absence of biology (Endler, 1973, Slatkin, 1985) - genetic description of organismal movement
- Implicit = active organism - emphasising behavs role = make active organism explicit part of evolutionary theory (Bateson, 1988; Ho, 1988; Wcislo, 1989)
- Behav traits distinct from other aspects of phenotype (Plotkin, 1988a) - foundation for arguments that behav plays unique role in evolution
- Due to reactive nature
- Doesn't mean all types behav and behav shifts affect evolutionary processes in same way
- Determine how diff types behavs and shifts affect selection pressures in ST to impact rate of evolutionary diversification

12

Fink (1982)

- model of Alberch et al. (1979) reformulated in terms phylogenetics - describe how heterochronic ontogenetic processes detected in nature
- heterochronic processes —> pedomorphosis —> morphologies which resemble primitive traits
- ontogenetic criterion for assessing polarity independent of hypotheses of phylogeny - as effective as outgroup comparison
- ontogenetic analysis aid detection of convergence but not in detection of parallelism

13

roots of comparative approaches and methods in behavioural and neuroscience research

- Focus on systematic comparisons by middle of 20th century
- ethology - heritage to school of thought
- experimental psych
- neuroscience
- Nobel prizes to physiologists and neurobiologists
- Lorenz et al. (1973)

14

ethology

- Is the "biological study of behaviour" (Tinbergen, 1963)
- emphasis on studying natural habitat and condition
- quantify the behaviours
- Is "the comparative study of behaviour which applies to the behaviour of animals and humans" (Lorenz, 1981)
- In the ethological approach animals are observed in their natural habitats or under conditions that are ecologically relevant
- Research focused on behaviour that has low interindividual variability - repeated behaviours
- locomotion (walking, swimming, flying)
- sequences (courtship, copulation, fights and contests, communication)
- fast responses (escpae, collision avoidance, capture)
- Good to quantify and compare with other species

15

ethology research

Radhakrishna and Sengupta (2020)

Jaffe et al. (2020)

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Radhakrishna and Sengupta (2020)

Although animals are the primary focus of studies in human-animal studies (HAS), very few ethologists engage with this discipline. Insights from HAS can help provide a deeper and richer understanding of animal behaviour and human-animal interactions. HAS perspectives regarding animal and human spaces, the sociozoological scale theory, and the concepts of animal agency and intersubjectivity in human-animal interactions help demystify puzzling aspects of human-wildlife conflict scenarios and impel a re-examination of ethological dictums and methodologies. We argue that inputs from human-animal studies will aid in the growth of ethology.

17

Jaffe et al. (2020)

Although animals are the primary focus of studies in human-animal studies (HAS), very few ethologists engage with this discipline. Insights from HAS can help provide a deeper and richer understanding of animal behaviour and human-animal interactions. HAS perspectives regarding animal and human spaces, the sociozoological scale theory, and the concepts of animal agency and intersubjectivity in human-animal interactions help demystify puzzling aspects of human-wildlife conflict scenarios and impel a re-examination of ethological dictums and methodologies. We argue that inputs from human-animal studies will aid in the growth of ethology.

18

behaviour is motor output

- Movement of different parts of the body (different muscle groups, appendages, limbs, vocal cords, syrinx)
- Decision-making in neuronal mechanisms to switch between movements and actions
- Variable and stereotyped elements of motor patterns
- Rapid responses and oscillatory activity (e.g. walking, running, flying - repeatedly switch muscle on and off)
- coordination and sequences of motor patterns
- Behaviour is measurable
- Link behaviour output to the motor output in the brain

19

classical ethological studies (Lorenz and Tinbergen, 1939)

- **Fixed acts** (elementary actions) and **fixed action patterns** (sequences of behavioural actions) are highly stereotyped behavioural responses
- e.g. speed of movement depending on heat of plate
- **Vacuum activity** - responses in absence of stimuli
- unrelated behaviour
- engage in sequence originally elicited by stimulus but also continued when stimulus no longer present
- Quantify behaviour and look at motor responses that happen in the brain

(see research doc)

20

classical ethological studies (Lorenz and Tinbergen, 1939) research

Berkenbeit et al. (1986)

Schleidt (1974)

Spena et al. (2020)

21

Berkenbeit et al. (1986)

Discusses the following factors underlying behavioral plasticity: (1) reflex adaptability and its role in voluntary movement control, (2) degrees of freedom and motor equivalence, and (3) the problem of the discrete organization of motor behavior. A variety of innate motor patterns are discussed, particularly the wiping reflex in the frog. It is proposed that central regulation of stretch reflex thresholds governs voluntary control over muscle force and length. This suggestion is an integral part of the equilibrium-point hypothesis, 2 versions of which are compared. It is concluded that voluntary movements are effected by the central nervous system (CNS) with the help of mechanisms that underlie the variability and modifiability of innate motor patterns.

22

Schleidt (1974)

- Within the concept of the “fixed action pattern” (FAP), as an elementary
unit of behavior, the fixedness (stereotypy, small variability) has become the most important diagnostic criterium. Few data are available, howcvcr, to
cvaluate the cxtent and the role of stereotypy of such behavior patterns and of their Constituent elements. Three kinds of stereotypy can be distinguished: (1) the degree of cotnpleteness with which the constituent elements of the pattern Are present, (2) the degree of coldpling between those elements in their concurrence or succession, and (3) the degree of variability of the various physical characteristics of the pattern as a whole, or of its constituent elements (duration, speed, position, intensity, etc.).
- Various methods, employed to investigate and measure stereotypy, especially in their degree of variability expressed by the coefficient of variation (V) or by the geometric standard deviation (s,,)~,), are described. They are exemplified by the gobbling call of the male turkey (Figs. 1 through 6, Tables 1 and 2) and by data from the literature (Figs. 7 and 8). Whether or not a particular behavior pattern appears to the un-aided eye as being stereotyped depends apparently on properties of the observer’s gestalt perception, which calculates a complex
impression of stereotypy from a variety of characteristics. No attempt has been made to express the overall stereotypy of a behavior pattern in a single index, but it is noted that whenever a characteristic of a behavior exhibits low variability (V 5 O.l), the behavior itself appears to the observer as being stereotyped.
- In FAPs of locomotion and related behaviors (e. g., preening, digging)
stereotypy seems related to their rhythmicity, which itself appears to be favored because of its efficiency. In FAPs of communicative function stereotypy is clearly related to the necessity for the transmitter and receiver to employ the same code, i.e., to be in tune. As a consequence, in a characteristic utilized for individual recognition intra-individual variability will be low, whereas the inter-individual variability of the same characteristic will be as high as necessary
to accomodate the individual code within the population.
- As a strategy for future research on elementary units of behavior it is
proposed that more attention be given to quantitative measurements of stereotypy than is currently being done and that the concept of fixed action pattern not be abandoned on the grounds that the term is incompatible with the notion of variability.

23

Spena et al. (2020)

Egg retrieval is a common behaviour among ground-nesting birds and a key part of their parental care. This paper deals with an experimental study of that behaviour in four Sicilian populations of Stone curlew, considering not only the success in retrieving but also the time latency in the bird reacting to the sight of an egg displaced from its nest and the relative retrieval time, which are two parameters that have been ignored until now by other studies. Egg displacements were made in 36 nests at the same distance of 50 cm. In each nest, two egg displacements were made, one in the first and the other in the second half of the brooding period, at a mean distance of 12.5 ± 3.5 days. In only seven out of 67 valid tests, the egg remained unretrieved, confirming the ability of Stone curlew to roll in their eggs. None of the factors considered as predictors had a significant effect on the success of the retrieval. Latency and retrieval times resulted positively correlated to each other, and were not statistically different among the four populations tested, nevertheless, showing a high variability even within the same population. In nine cases the retrieval was performed by the two parents acting in turn, however, in these cases the retrieval and not the latency times turned out to be longer with respect to the retrievals made by only one bird. When the retrieval times of the first displacement were compared to those of the second, the latter were shorter. The retrieval times are influenced by the ground conformation, being shorter in agricultural sites where the ground is plain and levelled. Indeed the GLMM analyses suggest a significant influence of different external factors on both the latency and retrieval times.

24

some behavioural patterns are stereotyped (fixed acts or fixed action patterns) can be triggered by specific stimuli (Tinbergen, 1952)

Some behavioural patterns are stereotyped (fixed acts or fixed action patterns) can be triggered by specific stimuli (Tinbergen (1952)

- Sign stimulus (releaser) and releasing values
- Testing retrieval behaviour with pairs of eggs - shape and size were critical sensory cues
- Male sickleback agression
- Greylag geese (Anser anser) - when an egg falls out of the nest as occasionally happens, the goose stretches out its neck and rolls its back with the underside of the bill moving its head from side to side as it goes to prevent the egg from rolling off the side. Sometimes maneouver fails and egg rolls away, but the goose continues as if it were still there. The same if egg is removed by experimenter

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some behavioural patterns are stereotyped (fixed acts or fixed action patterns) can be triggered by specific stimuli (Tinbergen, 1952) research

Fero et al. (2011)

Conte and Schulter (2013)

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Fero et al. (2011)

- Shortly after larval zebrafish become free swimming their behavior is modulated by both autochthonous signals and external stimuli. Larvae show rapid responses to a range of sensory cues but are also capable of executing extended behavioral programs in response to changes in the environment. At this early stage, larvae have a small repertoire of discrete stereotyped movements which are deployed in different contexts to generate appropriate behavior. We outline the range of behaviors defined in zebrafish larvae to date and discuss insights into neural function revealed by behavioral assays. A growing body of work demonstrates that tractability of behavior and neural connectivity in larval zebrafish facilitate the analysis of neural pathways underlying vertebrate motor control and sensory processing.

27

Conte and Schulter (2013)

Mate choice by phenotype matching, whereby individuals prefer a mate whose phenotype is similar to their own, should facilitate speciation with gene flow. This is because the genes that control mate signal (the phenotype being matched) also determine the preferred mate signal (“mate preference”). Speciation is made even easier if phenotype matching is based on a trait under
divergent natural selection. In this case, assortative mating should readily evolve as a byproduct of divergent selection on the trait. Previous observational studies of assortative mating between sympatric, hybridizing threespine stickleback species (Gasterosteus aculeatus complex) suggested that phenotype matching might occur by body size, a trait under divergent natural selection. To test this, we used experimental manipulation of body size to rule out the effects of confounding variables. We found that sizemanipulated benthic and limnetic stickleback females prefer mates whose body size more closely matches their own. It is thus likely that assortative mating by phenotype matching has facilitated the origin and persistence of benthic and limnetic threespine sticklebacks in the face of gene flow

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releaser stimuli can trigger behavioural output (Tinbergen and PErdeck, 1950; ten Cate et al., 2009)

- Pecks beak in order to be fed
- Cate - tested exposure and familiarisation - led to effect of red beak

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releaser stimuli can trigger behavioural output (Tinbergen and PErdeck, 1950; ten Cate et al., 2009) research

D'Eath (1998)

Bovet and Vauclair (2000)

Soffie and Zayan (1977)

30

D'Eath (1998)

The use of video images in place of natural stimuli in animal behaviour experiments is reviewed. Unlike most other artificial means of stimulus presentation, video stimuli can depict complex moving objects such as other animals, preserving the temporal and spatial patterns of movement precisely as well as colour and sounds for repeated playback. Computer editing can give flexibility and control over all elements of the stimulus. A variety of limitations of video image presentation are also considered. Televisions and video monitors are designed with human vision in mind, and some non‐human animals that differ in aspects of visual processing such as their colour vision, critical flicker‐fusion threshold, perception of depth and visual acuity, may perceive video images differently to ourselves. The failure of video stimuli to interact with subjects can be a drawback for some studies. For video to be useful, it is important to confirm that the subject animal responds to the image in a comparable way to the real stimulus, and the criteria used to assess this are discussed. Finally, the contribution made by video studies to date in the understanding of animal visual responses is considered, and recommendations as to the future uses of video are made