Biological psychology is the study of the relationship between psychological events and processes and physical events in the brain
Aims: To understand how the brain creates the mind and to uncover distinct psychological processes through their reflection in distinct brain processes
Issues: matching methods to the hypothesis being tested and ethical and practical considerations
Biological Psychology
Can discover how the brain works by either observing and measuring or manipulating
Biological and physiological method must be appropriate -> measure or manipulating
Matching methods to hypotheses
Causal or correlative
Species psychologically applicable or invasive
Spatio-temporal resolution physically applicable
Biological Psychology
Spatial and Temporal Scales
e.g., is the hypothesis about individual neurons or cerebral hemispheres
e.g., is the hypothesis about individual action potentials or long lasting changes in synapses
Biological Psychology - Relevant sizes (spatial)
Whole brain - 25 cm
Typical Topographic Map - 5 mm
Cortical column - 250 μm
Cortical layer - 50 μm
Neuron (varies) - 10 μm
Dendrite - 1 μm
Synapse - 100 nm
Biological Psychology - Relevant Times (Temporal)
Biological
Neural refractory period - 2ms
Signal time from eye to brain - 20-100ms
Psychological or behavioural
Time to make eye movement - ~100ms
Time to orient to spatial attention - ~150ms
Simple reaction time - ~250ms
Learning - seconds-minutes
Induced mood change - minutes
Lesions
Changes in psychological function accompanying brain damage may reveal something about the function of the damaged tissue
After accidental injury, damage, and abnormal change in humans, or intentional damage in animals
Phineas Gage - left frontal lobe highly damaged, leading to changes in personality to do with lacking inhibition and planning
Tan - impaired speech production, left frontal damage to 'Broca's Area' leading to 'Broca's Aphasia': limited speech and loss of grammatical structure. Later identification of Wernicke's Area: language comprehension
Lesion
HM - removal of parts of medial temporal lobes, including hippocampus, severe anterograde memory, inability to store and retrieve new memories, normal implicit learning: improved at tasks without memory of learning
Logic of Experimental Lesion Method
'Is brain region X important for task A' -> localisationist perspective
Ignoring adaptive and 'parallel' brain processes can lead to false conclusions
Lashley -> found learning occurred everywhere in the brain: larger lesion = larger deficit, harder task = more brain required -> 'principle of mass action'
Lesions - Single vs Double Dissociations
Single Dissociation - 1 lesion group, 1 control, 2 tasks
Believe lesion group to be impaired in one task but intact for other
Must assume both tasks are equally sensitive to group differences
Single dissociation may be from general effects of trauma
Dorsal lesions impaired at water maze task (spatial impairment)
Ventral lesions impaired at fear task (contextual fear impairment)
Involvement of hippocampus in contextual fear condition not a requirement for spatial learning
Deactivation Methods
Lesions destroy tissue permanently -> irreversible and invasive -> can't study humans
Reversibility allows animals to act as controls but can raise design issues
Deactivation - Chemical Methods
Drugs in small quantities to specific brain areas can deactivate specific regions temporarily -> i.e., muscimol for GABA
DREADDS - engineered receptors can be activated or deactivated by a certain drug allowing for changes over a sustained and short period of time
Deactivation Methods - Transcranial Magnetic Stimulation
An electrical pulse in a coil is used to induce a sudden change in a magnetic field in the area below it
This temporarily interferes with brain activity for a hundredth of a second
Good temporal resolution for cortical regions
Can have reasonable (<1cm) spatial resolution in conjunction with 3D MRI
No known side effects in humans
Neuroimaging Techniques - MRI
Manipulates the behaviour of hydrogen ions to yield radio signal - different types of tissue produce different radio signals
Maguire (2000) - posterior hippocampal volume correlated with time spent as a taxi driver -> spatial representation of environment and can expand to accommodate elaboration
Functional MRI - detecting blood oxygen level, a change in BOLD signal indicates a metabolically active brain region
Spatial resolution - 'voxels' - balance resolution with area covered and acquisition time, resolution can vary ~3mm
Temporal resolution poor
Neuroimaging Techniques - Functional MRI
Image Analysis: comparing signal strengths in different behavioural conditions, whole brain analysis involves tests on everyvoxel -> difficult due to many ways to correct for each test. Region of interest studies only test a few voxels per brain region - care needed picking the region
Criticisms
New phrenology - careful behavioural designs and statistical analysis
Over-emphasis of localisation of function
Correlation not causation
Early studies apply uncorrected statistical tests to neuroimaging data
Neuroimaging Techniques - Functional MRI - what it can tell a psychologist (Henson, 2005)
Function to structure inference - if different activity is detected under different task conditions, then the tasks are functionally dissociable
Structure to function inference - if the same pattern of activity is detected under two task conditions, then the tasks require a common function
Inferences rely on the assumption that there is localisation of function within the brain
Neuroimaging Techniques - EEG
Electrodes on the surface of the scalp record electroencephalograms
EEG activity is the results of electric fields generated by summing IPSPs or EPSPs in thousands or millions of cells
Spatial localisation possible with multiple electrodes
Multi-electrode EEG -> temporal resolution good, signals weak so need to be averaged over many trials, spatial resolution poor, poor for deep activity
Neural Recording - Electrophysiology: Single Cell Recording
Implanting a small electrode to pick up extracellular activity
Only picks up cells that are near and active at that time
O'Keefe (1975) - place cells that fire in the hippocampus in particular contexts/locations
High spatial and temporal resolution but small number of cells
High channel electrophysiology - thin probe rather than a bundle of electrodes -> hundreds of recording channels that can record thousands of cells in real-time
Neural Basis of Learning
Learning - change in behaviour resulting from experience
Looking for a physicalchange in the brain that corresponds with this learning
i.e., Pavlovian conditioning -> 2 stimuli become linked, generating a new behavioural response to a previously neutral stimulus
Learning is a psychological process of pairing events together - looking for a neural mechanism that would allow for associations to be formed
Neural Basis of Learning - Hippocampus
Important for learning and memory - cellular basis of learning and memory, processes multimodal sensory and spatial information
Kindling - idea that prior stimulation of neurons increases the likelihood of neurons firing
Pre-synaptic and post-synaptic populations must be segregated and easily identifiable - Bliss and Lorno (1973) - hippocampus
Stimulate the Perforant path, record post-synaptic activity in dentate gyrus
Some EPSP prior -> repeated stimulation -> Increased EPSP (long-term) -> weak stimulation now leads to strong response
Neural Basis of Learning - Long Term Potentiation
Key Properties: effects are long-term, and only occurs when firing of pre-synaptic neuron is followed by firing of post-synaptic neuron (co-occurrence)
Hebbian Basis of Memory - neurons that fire together, wire together
i.e., conditioning rabbit eye blinking -> strong sensory neuron and weak auditory neuron fires onto motor neuron -> repeatedpairing leads to strengthened response where tone alone is sufficient to generate a post-synaptic response -> motor neuron more receptive to weak auditory synapse
Neural Basis of Learning - Properties and Processes of Hippocampal LTP
NMDA receptor - 'fussy' - only opens in the presence of the neurotransmitter (glutamate) and when the post-synaptic membrane is depolarised -> pre- and post-synaptic activity coincidence detector
Ion-channel associated with the NMDA receptor is normally blocked by a positively charged magnesium ion -> depolarisation means the ion is no longer attracted
If pre-synaptic neuron continues to fire, glutamate continues to be released
Glutamate binds -> allows calcium to enter the cell
Tocco (1992) - LTP changes number of AMPA receptor numbers
AMPA is not 'fussy' -> only needs glutamate to bind to it
LTP requires AMPA receptors
Neural Basis of Learning - Hippocampal LTP: Post-Synaptic Changes 2: Protein Synthesis
Blocking protein synthesisprevents LTP (Nguyen, 1994)
The structure of the cell may change due to calcium entering the cell
Post-synaptic structural changes: long-term increase in AMPAR's, growth of new synapses, enlargement of synapses, splitting of synapses, enlarged post-synaptic area that glutamate can bind to
Neural Basis of Learning - Hippocampal LTP: Pre-synaptic effect
More radio-activity in synapse after LTP
Enhanced glutamate release in pre-synaptic cell -> interacts with increased area and receptivity -> easier to trigger
Retrograde transmitters from post-synaptic cell -> release of nitrous oxide that induces changes in the pre-synaptic cell -> production of glutamate is increased
Neural Basis of Learning - Long Term Potentiation Summary
LTP strengthens synaptic connections between neurons
Glutamate release activates AMPA receptors, frequent action potentials cause post-synaptic neuron to be depolarised
Magnesium ions no longer attracted to NMDA receptor -> calcium ions can flow in
Post-synaptic changes -> increases in AMPA receptors -> more receptive to glutamate
Post-synaptic cell releases NO (retrograde transmitters) to cause the pre-synaptic neuron to release moreglutamate
Neural Basis of Learning - LTP and learning and memory
Associative learning requires the formation of links between different stimuli, LTP is a mechanism for the formation of links between neurons
NMDA receptor (coincidence detector) -> LTP only occurs if cell is depolarised and glutamate binds to the receptor
Coincidence detection is important in associative learning -> i.e., neurons involved in 'food' are depolarised, and will only form synaptic connections with 'bell' neurons that are concurrently firing and releasing glutamate -> associations only formed with concurrent activity
Neural Basis of Learning - Is Hippocampal LTP required for Spatial Learning?
LTP observed in hippocampus and hippocampus involved in learning and memory, especially spatial learning
Morris water maze -> hippocampus-dependent
AP5 (NMDA receptor antagonist) failed to improve at task -> impaired spatial learning, but also showed sensorimotor impairment
Bannerman (1995) -> water mazes on separate floors
AP5impaired performance when there was no pre-training
AP5 did not impair performance when rats had pre-training
LTP in hippocampus may not be necessary for spatial learning
Motivation - Basic Drives: Homeostasis
The body strives to maintain a consistent environment -> biological needs impact behaviour, which is impacted by learning
When the body is challenged by an unavoidable loss of a regulated variable, there is an effect on motivation that leads to a change in behaviour
i.e., losing weight -> body has desire to maintain a certain weight -> restriction of food leads to a reduction in metabolic rate -> limiting extent of weight loss
Motivation - Basic Drives: Homeostasis - Brain systems for controlling hunger
Different regions of the hypothalamus have different roles in feeding behaviour
Lesions to the ventromedial hypothalamus result in overeating (hyperphagia) -> involved in satiety
Lesions to the lateral hypothalamus results in lack of hunger and weight loss (aphagia) - necessary for hunger
Motivation - Behaviour motivated by learning, not just immediate needs
Schepers and Bouton (2017) - rats trained to press a lever when either hungry or not
If behaviour is driven solely by needs, the hungry rat will press the lever more
Rats press the lever more in the sated condition -> context-dependent learning: learnt that when they are sated, food will appear -> not just current motivational state
Motivation - how motivational states affect behaviour
Hull's Drive Reduction Theory: drive - motivational state activated by a biological need; reinforcers satisfy needs and reduce drive; habit is what has been learnt about a reinforcer satisfying a need
Behavioural strength = Drive x Habit
If Drive or Habit = 0, there will be no behaviour
Bolles (1975) -> the more motivated (drive) a rat is (23h food deprivation), the more times they will press the lever; the more learning they have (reinforcement), the more times they will press the lever before extinction
Motivation - is Drive and Habit sufficient enough to describe adaptive behaviour?
Balleine (1992) -> pressing lever leads to food when either sated or hungry; sated or hungry during test
Hull assumes that motivational state during learning is not important and that you learn the 'habit' either way
Sated group -> increasing drive (hunger) did not increase behaviour -> have to learn that outcomes have a rewardingproperty
Hull: drive is a response to need, state of depletion motivates behaviours; evidence indicates behaviour is motivated by a desire to anticipate or avoid need
Learning and Motivation - Anticipating Need (Fitzsimons and Le Magnen, 1969)
Rats on a high protein diet need to drink a lot; when rats on a low protein diet switch to a high protein diet, they initially drink lots after meals
Eventually, rats increase their water intake prior to meals -> learnt association and anticipated future need for thirst
Learning and Motivation - Optimising Time and Effort (Collier and Johnson, 1997)
When it is easy to obtain food, animals will eat small amounts frequently
As it gets more costly and effortful to obtain food, meal size increases -> overall intake remains constant -> eating in an optimised way
Behaviour is structured in anticipation of needs and reduction of expenditure of effort
Learning and Motivation - Anticipation (Birch, 1991) - pre-school children
Children played in rooms where snacks were either available or not. Prior to test, children ate ice cream until full, then played in same room with snacks fully available
Children who played in room associated with snacks showed potentiated eating -> less time elapsed before they ate the snacks even when full; learning can lead to overeating
Association between cues and reinforcers can lead to a state of depletion being avoided
Learning and Motivation - Cue Potentiated Feeding (Johnson, 2013; Corbitt and Belleine, 2011)
Cues associated with food can lead to overeating when sated, even when drive is 0
Pavlovian to Instrumental Transfer: Stage 1 - Pavlovian, auditory cues; Stage 2 - Instrumental, lever pressing; Stage 3 - PIT, combined levers and audio cues
General PIT - behaviour usually increases -> response heightened if another cue shows that food will be available
Specific PIT - sound and lever associated with same reward leads to enhanced performance -> behaviours can be enhanced with cues to signal rewards
Learning and Motivation - Acquired Motivation
Rewards can also have an emotional effect that impacts behaviour
Contrast Effect: Negative Contrast -> Flaherty (1991) -> rats who went from a 32% to 4% sucrose solution drank less than rats who only ever drank 4%
Lick clusters show palatability -> as concentration increases, lick cluster size increases -> Austen and Sanderson (2016) -mice from 32% to 4% drank it like it was less palatable -> like they were disappointed
Learning and Motivation - Incentive Motivation (Spence)
Drive - motivation state driven by need; habit - the extent of learning; K - incentive motivation (motivating effect of reward) driven by prior experience
Behavioural Strength = Drive x Habit x K
i.e., Birch (1991) -> preschool children, during test drive was low but incentive motivation was high
i.e., Flaherty -> rats and sucrose, negative contrast effect reduced incentive motivation, even though drive was high
Learning and Motivation - Drive and Habit: conclusion
The need to maintain homeostasis results in motivational states that affect behaviour
Changes in behaviour are not just in response to biological needs, but also in anticipation of need
These anticipatory behaviours reflect learning of signals of reward and the incentivevalue of rewards
Learning and Motivation - Neural Basis of Reinforcement
Olds (1958) - rats with electrode in ventraltegmental area pressed lever to provide stimulation up to 700x per hour -> preferred lever pressing over food -> rewarding behaviour
Dopamine: stimulation of VTA leads to dopamine release, nucleus accumbens have dopaminergic neurons; also assesses value of stimuli, motor cortex leads to consequential behaviour
Nucleus accumbens - Mobbs (2003) - funny cartoons led to activation of the NAc -> integrating information and rewarding properties
Learning and Motivation - Dopamine and Reward
Wise (1978) - rats given Pimozide (dopamine receptor antagonist) stopped pressing lever to receive food -> like food was no longer rewarding
Anhedonia Hypothesis (Wise, 1982) -> good stopped tasting good; general reducedliking of stimuli -> dopamine = 'pleasure' chemical of brain?
Learning and Motivation - Dopamine and Liking
Parkinson's damages dopamine producing neurons, Seinkiewicz (2013) - patients did not differ in pleasantness rating of sweets
Non-verbal behaviour -> automatic reactions: hedonic vs aversive: Berridge and Robbinson (1998) - rats given 6-OHDA drug that selectively destroysdopaminergic neurons -> no reduction in hedonic reactions to sucrose -> no evidence that dopamine depletionreducesliking