Course Goals Neuroethlogy NROC34 • What is neuroethology? • Prof. A. Mason – Role of basic biology – Model systems (mainly invertebrate) • e-mail: – Highly specialized organisms – [email protected] – Biomimetics – [email protected] • Primary literature and the scientific process – No textbook – subject = NROC34 – But if you really want one, there are a couple of suggestion on the • Office Hours: Friday, 1 – 4:00 pm, SW566 syllabus • Basic principles of integrative neural function • Weekly Readings: download from course webpage – More on this a bit later www.utsc.utoronto.ca/amason/courses/coursepage/syllabus2012.html

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How? Evaluation • Case studies of several model systems – Different kinds of questions – Different kinds of research (techniques) • Weekly readings 5% – Sensory, motor, decision-making… • Mid-term 35-45% • Selected papers each week • Final Exam 45-60% – Read before, discuss during: usually readings are challenging and “discussion” means me explaining (so • Presentation (optional) 10% don’t be discouraged) • Most topics include current work • Weekly reading marks: each week several – Take you to the leading edge of research in selected people will be selected at random to areas contribute 3 questions about the papers.

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1 Information Neuroethology

• Who are you people? • The study of the neural mechanisms • What do you expect to learn in this course? underlying behaviour that is biologically • What previous course have you taken in: relevant to the animal performing it. – neuroscience • This encompasses many basic mechanisms – Behaviour of the nervous system. • Is this a req’d course for you? • Combines behavioural analysis and neurophysiology

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Neurobiology Behavioural Science

• What do I mean by “integrative neural function”? • What does the nervous system do? • Ethology • Psychology – detect information in the environment – biological behaviour – abstract concepts (e.g. – process the information to select appropriate behaviour – natural contexts learning, memory) – generate the appropriate behaviour – Lorenz, Tinbergen, – controlled conditions • How does the nervous system work? Von Frisch share • Why does it work that way? Nobel Prize 1973

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2 Environment Organism Senory Neural Motor input processing output physiology Activate Information from Integrate & muscles etc. to the environment analyse multiple Sensory Stimuli Adaptive Behaviour inputs create actions

nervous The flow of information through the nervous system system is the connection between stimulus and response.

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Behaviour • Early approaches – subjective, anecdotal, antrhopomorphic • Darwin – supplied a functional interpretation; did not have the immediate effect of stimulating more objective methods, but ultimately led Psychology, Ethology & Neurobiology in this direction. HISTORY – In the 20th century, application of objective methodology led to the development of scientific studies of animal behaviour in two major directions

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3 Pavlov’s Behaviour dog

• Psychology – Concentrated on a reductionist analysis in controlled (laboratory) conditions – Ivan Petrovich Pavlov and conditioned reflexes Two main ideas: Skinner Box – Burrhus Frederic Skinner and operant Learning conditioning Stimulus‐response

Skinner’s “Heir Conditioner”

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e.g. graylag goose egg-rolling Behaviour

Entire sequence always completed once • Ethology it starts…Fixed Action Pattern – Emphasized natural (or seminatural) condition and behaviours – Konrad Lorenz – Niko Tinbergen Any vaguely egg‐like object will – Concentrated on "innate" behaviours. Fixed do…Sign Stimulus (Innate action patterns, innate releasing mechanisms, Releasing Mechanism) action-specific energy, etc. Innate Behaviour: nature/nurture debate

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4 e.g. digger wasp homing Behaviorism behaviour • Habituation/dishabituation • Classical conditioning • Operant conditioning • Formation of new stimulus-response connections (or modification of existing ones) -- Learning

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Ethology

• Traditional ethology identifies “innate” or “natural” connections between stimuli and These are old-fashioned responses. concepts.

Behaviour is almost always much more complex.

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5 Back to the nervous system... Neurobiology • There are built-in mechanisms, activated by external stimuli, that generate the resulting • Traditionally disconnected from behaviour. – Anaesthetized animals (or pieces of them) behaviour. – Properties of small elements of the nervous system • Something changes as a result of experience • But there were some early efforts at inferring broader functions. that causes a modification of behaviour. – Charles Sherrington's Integrative Function of the Nervous System – emphasized reflexes as a fundamental functional unit • Both indicate there is something that could – nervous system acting to coordinate the action of many local be identified in the nervous system that reflexes (ie stimulus/response relationships) to guide to result is represents the physical basis of the overall goal-directedness of behaviour – Higher mental functions and emotions beyond the reach of behavioural phenomenon. mechanistic analyses.

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Neuroethology Other influences • Jakob von Uexküll – Umwelt – Interested in the animal’s “point of view” – Origins of the integrative view?

Just as morphological adaptations evolve to match an organism to its environment...

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6 Neuroethology Neurophysiology Review Relating nerve cells to behaviour (the super-simplified view) • Nervous system is composed of neurons that share certain basic features. • Basic features of neurons are common across all animals. …the structure and physiology of the • Electrical activity is the currency of the nervous system have nervous system. evolved to match behaviour and context.

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Neurophysiology Review Neurophysiology Review Measuring and manipulating Membrane potential electrical activity in neurons Semi-permeable membrane, intracellular recording opposing electrical and chemical gradients, amplifier -Volts+ electrode potential difference across membrane (inside negative re. outside) mV 0 -65

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7 Neurophysiology Review Neurophysiology Review Action potential Synaptic transmission

Electrical impulse that propogates along a neuron

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Neurophysiology Review Function of the nervous system Excitatory = depends on... depolarizing; tending to make post-synaptic cell fire an action potential • Connectivity (interconnections between neurons) Inhibitory = • Intrinsic properties of neurons hyperpolarizing; tending to stop post- synaptic cell from firing an action potential

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8 Connectivity Connectivity

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Connectivity Connectivity

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9 Modulation & Plasticity Intrinsic Properties

Tonic

Phasic

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Identifying the neural basis of Behaviour + Neurophysiology behaviour • Correlation • “Natural” behaviour – what neural activity is reliably associated with - What the nervous system is “designed” to particular stimuli and/or responses? produce • Necessity – if neural activity is suppressed, is behaviour also suppressed? • Sufficiency – if neural activity is generated artificially, does it result in behaviour?

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10 e.g. crayfish tailflip Crayfish tail-flip circuitry

Correlation

Sufficiency

Necessity

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Neural Circuitry

Senory Neural Motor Neural input processing output processing

Information from Integrate & Activate the environment analyse multiple muscles etc. to inputs create actions Motor output Senory input The flow of information through the nervous system Integrate & Activate is the connection between stimulus and response. Information from the environment analyse multiple muscles etc. to inputs create actions

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11 August Krogh Invertebrates

• Krogh’s Principle: “For a large number of problems there will be some animal of choice on which it can be most conveniently studied.”

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Distributed, ventral CNS Invertebrate nervous systems

• Ventral • Distributed, ganglia and connectives • Large neurons • Relatively simple circuits (“identified neurons”) • Very robust

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12 Identified Neurons Large Neurons

N. Hatsopoulos, F. Gabbiani and G. Laurent (1995). Elementary Computation of ObjectApproach by a Wide Field Visual Neuron, Science,270:1000-1003. NROC34 2012:1 49 NROC34 2012:1 50

Very robust

• Detailed analyses • Fundamental mechanisms – neurophysiology – neuroethology - link between behaviour and underlying mechanism • Understanding and application...

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13 Fundamental Mechanisms Caenorhabditis elegans

959 cells (302 neurons)

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Applications

It’s not only invertebrates

Toad Frog

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14 Topics

• Hearing – Cricket phonotaxis • Vision – Bee navigation – Fly motion detection • Olfaction – Locust smell processing • Pattern generation – Neural networks •Others? – Electroreception – Echolocation – Robotics

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