Midterm 2 Notes

February 13, 2024 8:53 PM

Chapter 3:
5. Biological Basis of Behaviour

Nervous System Overview:

Divided into two parts:
CNS:
Brain, Spinal Cord, Optic Nerve, Retina

PNS:
Cranial Nerves, Spinal Nerves, Autonomic Nervous System: Sympathetic NS, Parasympathetic NS, and Enteric NS, Somatic Nervous System, etc.

The Brain:
Neurons:
The basic unit of the nervous system, each composed of a cell body, dendrite, and axon.
Human brains contain approx. 67-86 billion

Glial Cells:
Nonneuronal brain cells that provide structural, nutritional, and other types of support to the brain.
Also called neuroglia.
4 different types of glial cell
Approx. 40-50 billion

The Parts of a Neuron:

Soma (Cell Body):
May have, one, two, or many dendrites and typically one axon.
Synthesize macromolecules, integrate electrical signals

Dendrites:
Tapered extensions of cells body
Collect information from other neurons

Axon:
Single, cylindrical; may be many centimeters long; may be myelinated or unmyelinated
Conduct information to other neurons.
Sometimes has a Myelin Sheath which allows for faster transportation of information/signal

Axon Terminals (presynaptic terminals/synaptic boutons):

Vesicle-filled apposition to part of another neuron
Transmit information to other neurons

The Axon Terminal:

Synaptic Vesicle:
Spherical sac containing neurotransmitters.

Neurotransmitter:
The chemical released from the (presynaptic) axon terminal that serves as the basis for communication between neurons.

Synapse:
An intercellular site where fast, highly localized transmission of chemical and electrical signals occur.
Includes the pre-synaptic and post-synaptic terminals as well as the synaptic cleft.

Synaptic Cleft:
A gap into which neurotransmitters are released from the axon terminal.

Types of Neuron:
Unipolar Neuron and Bipolar Neuron:

Multipolar Neurons:
Multipolar Neurons:

The Neural Impulse:

Resting Potential:
Difference in electrical charge (-70 millivolts) across the neuronal membrane, when the neuron is not being stimulated or inhibited.

Depolarization:
The reduction of a membrane's resting potential so that it becomes less negative.

Action Potential:
Electrical impulse that travels down the axon triggering the release of neurotransmitters.

Repolarization:
Na+ channels close, K+ channels open
K+ efflux down electro-chemical gradient

Hyperpolarization:
"Overshoot" of repolarization.

Refractory Period:
The time during which another action potential is impossible; limits the maximal firing rate of the neuron.
Absolute
Relative

Receiving Signals:
Postsynaptic Potential (PSP):
A voltage change at a receptor site on a postsynaptic cell membrane.

Excitatory PSP:
A positive (+) voltage shift that increases the likelihood that the postsynaptic neuron will fire action potentials (because it gets closer to the -55 mV threshold).

Inhibitory PSP:
A negative (-) voltage shift that decreases the likelihood that the postsynaptic neuron will fire action potentials (because it gets further from the -55 mV threshold).

Reuptake:
Reabsorption of neurotransmitters from the synaptic cleft by the presynaptic membrane.
Neurotransmitters:

Small-Molecule Neurotransmitters:
Amino Acids: Glutamate, Aspartate, Glycine, GABA
Monoamines:
2 types:
Catecholamines: Dopamine, Epinephrine, Norepinephrine
Indolamines: Serotonin
Acetylcholine: Acetylcholine
Unconventional Neurotransmitters:
2 types:
Soluble gases: Nitric oxide, Carbon monoxide
Endocannabinoids: Anandamide

Large-Molecule Neurotransmitters:
Neuropeptides: Pituitary peptides, Hypothalamic peptides, Brain-gut peptides, Opioid peptides, Miscellaneous peptides, Endorphins

Acetylcholine (ACh):
Released by motor neurons controlling skeletal muscles.
Contributes to the regulation of attention, arousal, and memory.
Nicotine stimulates ACh receptors.
Memory enhances increase ACh.
Insecticides block the breakdown of ACh creating excess amounts.
Botox causes paralysis by blocked ACh release.
Alzheimer's disease is associated with low levels of ACh.
Agonist:
A chemical that mimics the action of a neurotransmitter when it binds with the neurotransmitters receptor site.
Antagonist:
A chemical that blocks the action of a neurotransmitter when it binds with the neurotransmitters receptor site.

Monoamines:
Dopamine (DA):
Contributes to control of voluntary movement
Cocaine and amphetamines elevate activity at DA synapses
Dopamine circuits in medical forebrain bundle characterized as "reward pathway"
Degeneration of DA neurons is believed to be responsible for symptoms of Parkinson's disease.

Serotonin (5-HT):
Involved in regulation of sleep and wakefulness, eating, aggression.
Prozac and similar antidepressant drugs affect serotonin circuits.

Norepinephrine (NE):
Brain arousal and other functions like mood, hunger, sleep, and sexual behaviour.
Drugs such as amphetamines and methamphetamines increase NE levels.
Also known as noradrenaline.
Functions as both a hormone and a neurotransmitter.

Amino Acids:
GABA (Gamma-aminobutyric acid):
The brain's main inhibitory transmitter
Contributes to regulation of anxiety and sleep/arousal
Valium and similar antianxiety drugs work at GABA synapses.

Glutamate:
Main excitatory neurotransmitter in the nervous system; participates in relay of sensory information and learning.

Unconventional Neurotransmitters:
Anandamide:
Binds to the same receptors as THC (tetrahydrocannabinol), the active ingredient in cannabis leaves.
Comes from the Sanskrit word "Ananda", which means "joy, bliss, delight".
Occurs in both the CNS and PNS.
Likely plays a role in eating, memory, motivation, and sleep.
Pain, fear, and healing? (decreased pain, fear, but increased healing)

Neuropeptides:
Endorphins:
A naturally occurring (endogenous) analgesic (painkiller).
Inhibit the communication of pain signals to the spinal cord.
Insensitivity to pain (oversupply)
Hypersensitivity to pain (undersupply).

The Endocrine System:

Series of glands that produce chemical substances known as hormones.
Hormones:
"Slow" chemical messenger released into the blood by endocrine glands.

Neural Plasticity (Neuroplasticity):

Plasticity:
Neural Plasticity in Early Development:
1. Growth of dendrites and axons
2. Synaptogenesis - Formation of new synapses.
3. Pruning - The removal of extra synapses to increase the efficiency of a neural network. Occurs until about age 10. Approx. 70% of neurons die off.
4. Myelination - The insulation of axons with a myelin sheath.

Neural Plasticity and Learning:

Long-Term Potentiation of synapses:
A long-lasting enhancement in signal transmission between two neurons that result from stimulating them synchronously.

Neural Plasticity Following Injury:

Brain regions can sometimes take over functions previously performed by others.
Researchers are searching for treatments to promote healing and prevent damage.
Stem cell:
A cell (often originating in embryos) which has the capacity to differentiate into a specialized cell.
Potential to offer treatments for Alzheimer's Stroke, Parkinson's, Diabetes, Bindless, Deafness, Infertility, and numerous other things
Neurogenesis:
Creation of new neurons in the adult brain.

The CNS:
Meninges:
Three protective membranes that cover the brain and spinal chord.
Dura mater, Arachnoid mater, Pia mater

Cerebral Ventricles:
Pockets in the brain that contain cerebrospinal fluid (CSF), which provide the brain with nutrients and cushion against injury.
Note: CSF is also found in the subarachnoid space.

Franz Joseph Gall (1758 - 1828):

All mental functions arise from the brain.
Mind and body are NOT separate entities.
Brain consists of functional regions.
Also a proponent of phrenology unfortunately.

The Cerebral Cortex:

Cerebral Cortex:
Outermost part of forebrain, responsible for analyzing sensory processing and "higher" brain functions.

Cerebral Hemispheres:
Two halves of the cerebral cortex, each of which serve distinct yet highly integrated functions.

Corpus Callosum:
Large band of fibers connecting the cerebral hemispheres:

The Frontal Lobe:

Performs functions that coordinate other brain areas, motor planning, language, and memory.
Primary Motor Cortex: Part of the frontal responsible for bodily movement.
Prefrontal Cortex: Part of the frontal lobe responsible for thinking, planning, and language.
Broca's Area: Language area in the prefrontal cortex that helps to control speech production and some aspects of speech comprehension.

The Parietal Lobe:

Processes touch information, integrates vision and touch.
Primary Somatic Sensory Cortex (somatosensory cortex): Regions of the cerebral cortex that initially process information from the senses.

The Temporal Lobe:

Processes auditory information, language and autobiographical memory.
Wernicke's Area: Part of the temporal lobe involved in understanding speech.

The Occipital Lobe:

Back part of cerebral cortex specialized for vision.

The Basal Ganglia:

A group of nuclei (clusters of neurons) located beneath the cerebral cortex.
Involved in goal-directed motor control.
Contains dopamine neurons and is closely associated with reward and motivation.

The Limbic System:

A loosely connected network of structures located roughly along the border between the cerebral cortex and deeper subcortical areas.
Plays a role in olfaction (smell), Motivation, Memory, and Emotion

The Thalamus:
Gateway from the sense organs to the primary sensory cortex.
All sensory information (except smell) is relayed through the thalamus.

The Hypothalamus:
The part of the brain that regulates the pituitary gland and is responsible for maintaining a constant internal state.
The four F's: Fleeing, Fighting, Feeding, Sexual Behaviour (Fornication)
Body temp
Hunger/thirst
Sleep
Emotional behaviours

The Hippocampus:
Part of the brain that plays a role in spatial memory and may be necessary for the formation of new memories.

Amygdala:
Part of limbic system that play key roles in fear, excitement, and arousal.

The Brain Stem:

Part of the brain between the spinal cord and cerebral cortex that contains the midbrain, pons, and medulla.

Midbrain:
Part of the brain stem that contributes to movement, tracking of visual stimuli, and reflexes triggered by sound.

Hindbrain:
Pons, medulla, and cerebellum.

Medulla:
Part of brain stem involved in basic functions, such as heartbeat and breathing.

Pons:
Part of the brain stem that connects the cortex with the cerebellum and relays information between the two.

Cerebellum:
Hindbrain structure responsible for smoothing and finessing fine motor control initiated by other brain regions.
Likely plays a role in other areas: Executive function, Spatial abilities, Aspects of language

Reticular Formation:
A complex network of about 100 tiny nuclei that occupies the central core of the brain stem.
Contains the Reticular Activating System:
The hypothetical arousal system in the reticular formation.
Responsible for arousal and consciousness.

The Spinal Cord and the Reflex Arc:

Spinal Cord:
Thick bundle of nerves that conveys signals between the brain and the body.

Interneuron:
Neuron that sends messages to other neurons nearby.

Reflex:
An automatic motor response to a sensory stimulus.

The PNS:
The part of the nervous system that lies outside the CNS.
Divided into two main sections:
The Somatic NS:
Part of the nervous system that conveys information between the CNS and the body, controlling and coordinating voluntary movement.
The Autonomic NS:
Part of the nervous system controlling the involuntary actions of our internal organs and glands, which (along with the limbic system) participates in emotion regulation).
Divides into two more main sections:
Sympathetic NS:
Division of the ANS engaged during a crisis or after actions requiring fight or flight.
Parasympathetic NS:
Division of ANS that controls rest and digestion

Measuring the Brain:

Phrenology:
Basically judging the size and shape of your head to what intellect and personality you have.

Brain Damage:
Localized brain functions can be assessed by examining brain damage and seeing what behavioural issues arise from it
Via post-mortem examination or stereotaxic lesions in lab animals.
Measurement via devices like magnetic resonance imaging (MRI)
Neuropsychological testing

Electroencephalograph (EEG):
Records the brain's electrical activity at the surface of the skull.

Pros:
Used in human and non-human animals.
Detects rapid changes in electrical activity (high temporal resolution)
Non-invasive

Cons:
Shows only large aggregates of neural activity (has low spatial resolution).

Non-functional Brain Scans (neuroimaging):

Computed Tomography (CT or CAT scan):
A scanning technique using multiple X-rays to construct 3D images.

Magnetic Resonance Imaging (MRI):

Technique that uses strong magnetic fields to indirectly visualize brain structure.
Better suited than CT scans for visualizing soft-tissues like brain tumors.

Functional Brain Scans:

Positron emission tomography (PET):
An invasive imaging technique that measures consumption of glucose-like molecules, yielding a picture of neural activity in different regions of the brain.

Functional MRI (fMRI):

Technique that uses magnetic fields to visualize brain activity using the BOLD response.
BOLD = Blood Oxygenation level dependent.
Highly sensitive to motion
Questions about data analysis

Magnetic Stimulation:
Transcranial Magnetic Stimulation (TMS):
A technique that applies strong and quickly magnetic fields to the surface of the skull that can either enhance or interrupt brain function.
Non-invasive

Neuroanatomical Techniques:
Golgi Stain:
A neural stain that completely darkens a few of the neurons in each slice of tissue, thereby revealing their silhouettes.

Nissl Stain:
A neural stain that has an affinity for structures in neural cell bodies.

Electron Microscopy:
A microscopy technique used to study the fine details of cellular structure.

Genes, Evolution, and Behaviour:

Genetic Expression:
Chromosome:
Threadlike structures inside a cell's nucleus that carries genes.

Gene:
The section of a chromosome that controls the synthesis of one protein.
Composed of DNA

Genome:
Set of all types of genes distinguished by their locus and base sequence.

Allele:
Any one of two or more genes that may occur alternatively at a given site on a chromosome.

Genotype:
An organism's genetic makeup.
Phenotype:
An organism's observable traits.

Dominant Gene:
A gene that mask's the effect of other genes.

Recessive Gene:
A gene that is expressed only in the absence of a dominant gene.

Polygenic Transmission:
When a single phenotypic trait is controlled by more than one pair of genes.

Epigenetics:
The study of heritable changes in gene function that do not involve changes in DNA sequence.

Genetic Engineering:
Knockout Procedure:
A technique in which a gene is made inoperative.
Behaviour of KO'd animal is compared to a normal animal.
More than one gene can be KO'd.
Genes can also be "knocked-in" (replaced).
Problems: Behaviour is often the product of multiple genes (not one), A single gene can have multiple effects.

Behavioural Genetics:
Heritability:
The extent to which the variation of a particular trait across individuals of a particular population is due to genes.
Estimated using the heritability coefficient.
Cannot be used to make statements about individuals.
Can only be applied within a single group.

Heredity:
The passing of traits from parents to offspring by way of genes.

Concordance:
The likelihood that two people share the same characteristic.

Designs in Behavioural Genetics:

Family studies
Twin studies
Adoption studies

Family Studies:
Analysis of how characteristics run in intact families.
Useful for estimating risk of a disorder among the relatives of members afflicted with the disorder.
Problems: Relatives share similar environment and genes (can't separate genes and environment).

Twin Studies:
Twins can be of two types:
Identical:
Come from single zygote that split (monozygotic; MZ).
Share 100% of genes.
Fraternal:
Come from two separate zygotes (dizygotic, DZ).
Share 50% of genes.
Logic: If MZ twins show more similarity on a particular trait than DZ twins, this is assumed to be reflective of genetic factors.
Within both types of twins, environment is assumed to be controlled: Both types of twins grow up in the same home, at the same time, have similar neighbors, education, etc.

Adoption Studies:
An analysis of how traits vary in individuals raised apart from their biological relatives.
Adopted Person - Shares genes and not environment with biological parents and vice versa with adopted parents.
Logic:
If the adopted person shows a greater similarity to biological parents on a psychological trait, the trait is assumed to be more heavily influenced by genes.
If the adopted person shows a greater similarity to adopted parents on a psychological trait, the trait is assumed to be more heavily influenced by environment.
Selective Placement Confound:
Adoption agencies try and place children in homes similar to that of their biological parents.
Adoption studies assume that placement of the child in a home is random and not based on socioeconomic or psychological status of the biological family.

Reaction Range:
The genetically influenced limits within which environmental factors can exert their effects on an organism.

The Evolutionary Basis of Behaviour:

Evolutionary Psychology:
A discipline that applies Darwin's theory of Evolution by Natural Selection to human and non-human animal behaviour.
Criticisms:
Many claims of Evolutionary Psych are not testable and/or unfalsifiable (behaviour leaves no fossil record).
Relies on many assumptions about early humans which we know very little about.
Multiple "evolutionary" explanations are often possible and can confirm to any finding we like.

Chapter 4:
6. Sensation and Perception (review pages 137 - 143, 144 - 148, 148 - 154)

Sensation:
The process by which (objective) stimuli are detected, transduced into nerve impulses, and sent to the brain.

Perception:
The brain's interpretation of raw sensory inputs.

Sensation:
Transduction:
The process of converting an external energy or substance into electrical activity within neurons.

Sense Receptor:
Specialized cell responsible for converting external stimuli into neural activity for a specific sensory system.

Sensory Adaptation (neural adaptation):

Activation is greatest when a stimulus is first detected and then declines in responsiveness over time.

Psychophysics:
The study of how we perceive sensory stimuli based on their physical characteristics.
Absolute Threshold:
The lowest level of a stimulus needed for the nervous system to detect a change 50% of the time.
Just Noticeable Difference (JND):
The smallest change in the intensity of a stimulus that we can detect.
Weber's Law:
There is a constant proportional relationship between the JND and original stimulus intensity.
I = Intensity of the stimulus
K = Constant
JND = K * I

Signal Detection Theory:

Theory regarding how stimuli are detected under different conditions.
Accuracy = Number of Correct Responses/Number of Attempts
We use 2 questions - Is the Stimulus Present? and Did you Perceive the Stimulus?
Signal-to-noise ratio:
The ratio of the power of a signal to the power of background noise.
Signal = Stimulus to be perceived
Noise = Everything else
SNR = P_signal/P_noise
Increased sensitivity to the signal causes hits and correct rejections to occur more often.
Decreased sensitivity to the signal causes misses and false alarms to occur more often.
Differences in sensitivity are measured with a statistic called d' (d-prime).
A measure of the stimulus' salience.
Increases in d' represent improved detection.
Plotted using ROC curves (Receiver Operating Characteristic Curves)

Polygraphs:
Accuracy depends on:
Whether the person is lying
Examiner's set thresholds
Population the person belongs to.
Circumstances of the test
Test conditions
Whether the subject employs countermeasures
Legal and institutional policies
Relationship/interaction with the examiner
Subject's expectation/understanding about the test
Etc.

Examiner's set thresholds need to be based on all these factors.

Perception:
Perceptions is determined by three basic elements:
What is currently in the sensory field.
What was in the sensory field a moment ago.
What we have experienced in the past.
Parallel Processing:
The ability to attend to many sense modalities simultaneously.
Bottom-Up Processing:
Processing in which a whole is constructed from parts.
Top-Down Processing:
Conceptually driven processing influenced by beliefs and prior learning.

Bottom-up and Top-down processing are not mutually exclusive.

Perceptual Set:
A set formed when expectations influence perceptions.

Perceptual Constancy:
The process by which we perceive stimuli consistently across varied conditions.

A disclaimer:
Perceptual sets and constancy are just concepts which describe patterns of behaviour in relation to sets of stimuli.
They do not provide an explanation for that pattern of behaviour.

Attention:
Selective Attention:
The process of selecting one sensory channel and ignoring or minimizing others.
Assumed to be controlled by the reticular activating system (RAS) and higher cortical regions.
Filter 'Theory' of Attention:
Attention is a "bottle neck" through which information passes.
Dichotic Listening: a research design in which subjects have a message delivered to each ear independently through headphones. Both messages are delivered simultaneously,
but only the attended ear's message is able to be accurately recalled.

Inattentional Blindness:
Failure to detect stimuli that are in plain sight when our attention is focused elsewhere.
Change Blindness:
Failure to detect changes in a visual stimulus.

Subliminal Messaging

Extrasensory Perception (ESP):

Perception of events outside the known channels of sensation.
Precognition:
Prediction events before they occur through paranormal means (don't rely on standard scientific mechanisms).
Telepathy:
Reading other people's minds.
Clairvoyance:
Detecting the presence of objects or people hidden from view.

Why do people believe in ESP?

Illusory Correlations
People underestimate the frequency of coincidences.
The Birthday Problem:
How large must a group of people be before the probability of two people sharing the same birthday exceeds 50%?
It's about 23

The Visual System (real sensory perception):

Visible Light:
Electromagnetic radiation between 400 and 700 nm.
Hue:
The colour of light.

The Eyeball:
The Cornea:
Part of the eye containing transparent cells that focus light on the retina.
Myopia (nearsightedness):
Cornea is too long causing the focus of light to occur in front of the retina.
Can only see close objects well.
Hyperopia (farsightedness):
Cornea is too flat causing the focus of light to occur behind the retina.
Can only see far objects well.

The Lens:
Part of the eye that changes curvature to keep images in focus.
Contains transparent cells (light passes through them).
Accommodation:
Changing the shape of the lens to focus on objects near or far.

The Retina:
Membrane at the back of the eye responsible for converting light into neural activity.
Rods:
Photoreceptor cells in the retina allowing us to see in low levels of light.
Approx. 92 million per eye.
Dark Adaptation: Time in dark before rods regain maximum light sensitivity.
Cones:
Photoreceptor cells in the retina allowing us to see in colour.
Approx. 6-7 million per eye.
Photopigments:
Protein molecules within the rods and cones whose chemical reactions when absorbing light result in nerve impulses being generated.
Optic Nerve:
Nerve (technically a bundle of ganglion cell's axons) that travels from the retina to the brain.

Blind Spot:
Part of the visual field we can't see because of an absence of rods and cones (optic nerve's location).

Visual Perception:

Shape and Contour:

Hubel and Wiesel:
Recorded electrical activity of V1 area of visual cortex in cats.
Discovered feature detector cells:
Cells that detect lines and edges.
At later levels of visual processing (V2) cells begin to detect more complex shapes and movements.

The Principles of Gestalt Psychology:

A German school of psychology that emphasized the natural organization of perceptual elements into wholes or patterns.
"Gestalt" = "whole"
Psychology should be focused on how the "whole" is created from its basic parts.
Main Figures: Max Wertheimer, Wolfgang Kohler, Kurt Koffka
Proximity:
Objects physically close to each other tend to be perceived as unified wholes.
Similarity:
Similar objects will be seen as being grouped together or related.
Continuity:
We perceive lines as continuous movement while discounting abrupt changes.
Also called "good continuation"
Closure:
The tendency to view incomplete figures or forms as complete objects.
Symmetry:
Symmetrically arranged are perceived as wholes.
Figure-Ground Segregation:
The tendency to separate elements of an image into a foreground (figure) and background (ground).

Colour:

Trichromatic Theory:
Idea that colour vision is based on our sensitivity to the three primary colours.
Fits with later findings that three types of cone cells are present that respond maximally at different wavelengths.
Short, Medium, and Long
Relies on additive colour mixing.
Can't account for afterimages.

Opponent Process Theory:

Theory that we perceive colors in terms of three pairs of opponent colors:
Red or green
Blue or yellow
Black or white
Dual-Process Theory:
The modern colour vision theory that posits that cones are sensitive to red, green, and blue as well as opponent processes at the level of ganglion cells.

Colour Blindness:
Inability to see some or all colours.
Usually due to genetic abnormalities that cause the absence/reduction in a specific cone type.
Monochromats - have only one type of cone (very rare = 0.0007% of the population).
Dichromats - have two types of cone.
Brain damage to cortical areas responsible for vision can also produce colour blindness.

Depth Perception:
Ability to judge distance and 3D relations.

Monocular Cues:
Stimuli that enable us to judge depth using only one eye.
Relative Size: All things being equal, more distant objects look smaller than closer objects.
Texture Gradient: The texture of objects become less apparent as objects move farther away.
Interposition: Close objects block the view of distant objects.
Linear Perspective: The outlines of objects converge as distance increases.
Height in plane: Here distant objects tend to appear higher, and nearer objects appear lower.
Light and Shadow: The casting of shadows give objects a 3D form.
Motion Parallax: When objects moving across a field of view appear to move more/faster the closer they are to an observer.

Binocular Cues:
Stimuli that enable us to judge depth using both eyes.
Binocular Disparity: Depth information is obtained by comparing the difference in image location of the left and right eyes.
Binocular Convergence: The degree to which your eyes converge inward gives provides information to estimate distance.

Blindness (Vision Loss):

The inability to see due to problems with the eye and its related structures.
Vision is less than or equal to 20/200 (20/20 is considered normal vision).
Rely more on other senses (touch, hearing, etc.).
Visual cortex's function changes due to neural plasticity.

Motion Blindness:
A neurological disorder in which a person is not able to perceive motion.
Also known as "Cerebral Akinetopsia"
Often caused by: Brain damage, Alzheimer's disease

Visual Agnosia:
A failure to recognize visually presented objects.
Not due to impairments in memory or intelligence.
Often due to damage to peripheral regions of occipital cortex near the parietal and temporal lobes.
Different types:
Prosopagnosia: failure to recognize faces.
Anosognosia: failure to recognize illness.
Finger agnosia: failure to recognize fingers.
Topographical disorientation: failure to recognize places.

Blindsight:
The ability of individuals with blindness to detect and respond to visual stimuli despite lacking awareness of having seen anything.
Often the result of damage to V1 area of visual cortex.

From Textbook Readings for Chapter 4:

1. Hearing

The sense of hearing depends on sound waves, changes in air pressure unfolding over time. Just as vision is about the perception of meaningful visual objects, hearing involves
transforming changes in air pressure into meaningful sound objects (or sources).
There are three physical dimensions of a sound wave that correspond to dimensions of auditory perception: frequency, amplitude, and complexity.

The frequency of the sound wave depends on how often the peak in air pressure passes the ear or a microphone, measured in cycles per second, or Hz. The repetition rate of a
sound wave is perceived as the pitch, how high or low a sound is.

The amplitude of a sound wave refers to its intensity, relative to the threshold for human hearing (set to 0 dB). Loudness is the perception of a sound's intensity. Sounds above 85
dB can cause hearing damage, depending on the length of exposure.

The complexity of sound waves, or the mixture of frequencies, influences perception of timbre, the quality of sound that allows you to distinguish two sources with the same
pitch and loudness.

The human ear is divided into three distinct parts:

The outer ear collects sound waves and funnels them towards the middle ear, which transmits the vibrations to the inner ear, embedded in the skull, where they are transduced
into neural impulses.

The outer ear consists of the visible part on the outside of the head (called the pinna); the auditory canal; and the eardrum, an airtight flap of skin that vibrates in response to
sound waves gathered by the pinna and channelled into the canal. The middle ear, a tiny, air-filled chamber behind the eardrum, contains the three smallest bones in the body,
called ossicles. Named for their appearance as hammer, anvil, and stirrup, the ossicles fit together into a lever that mechanically transmits and amplifies vibrations from the
eardrum to the inner ear. Amplification is required because the ossicles push against the oval window, which is a membrane that separates the middle ear from the cochlea of
the inner ear. The ossicles take the airborne pressure wave at the eardrum and transfer it into a pressure wave in fluid. Fluid requires more energy to vibrate.

The inner ear contains the spiral-shaped cochlea, a fluid-filled tube that contains cells that transduce sound vibrations into neural impulses. The cochlea is divided along its length
by the basilar membrane, a structure in the inner ear that moves up and down in time with vibrations relayed from the ossicles, transmitted through the oval window. Sound
causes the basilar membrane to move up and down in a travelling wave. The frequency of the stimulating sound determines where on the basilar membrane the up-and-down
motion is highest. When the frequency is low, the wide, floppy tip (apex) of the basilar membrane moves the most; when the frequency is high, the narrow, stiff end closest to
the oval window (base) moves the most.

When the basilar membrane moves up and down, the cochlear fluid moves back and forth. This stimulates thousands of inner hair cells, which are specialized auditory receptor
neurons embedded in the basilar membrane. The hair cells have long hairs sticking out their tops that bend back and forth in the cochlear fluid, like seaweed in a current. This
back-and-forth bending generates rhythmic action potentials in the auditory nerve axons that travel to the brain, and these action potentials are precisely timed with the
pressure peaks of the original sound wave. The auditory nerve axons that fire the most are those connected to hair cells in the area of the basilar membrane that moves the
most.

From the inner ear, action potentials in the auditory nerve travel to several regions of the brainstem in turn, then to the thalamus and ultimately to an area of the cerebral cortex
called area A1, the primary auditory cortex in the temporal lobe. There is some evidence that the auditory cortex is composed of two distinct streams, roughly analogous to the
dorsal and ventral streams of the visual system. Spatial ("where") auditory features, which allow you to locate the source of a sound in space, are handled by areas towards the
back of the temporal lobe and in regions that may overlap with the visual dorsal stream. Features that allow you to identify the sound (what it is) are handled by areas in the
lower (ventral) part of the temporal lobe, and this pathway may overlap with the ventral visual pathway.

Your brain uses patterns of action potentials in the auditory nerve to perceive sounds in terms of their loudness, pitch, timbre, and location. How does it accomplish this?
 • The perceptual attribute of loudness is signaled by the total amount of activity in hair cells, rather like how brightness of light is signaled by the total amount of activity in
photoreceptors.
• How we perceive pitch has been studied extensively, and it seems to depend on two aspects of auditory nerve activity. First, different frequencies stimulate neural signals
at specific places along the basilar membrane. This provides a place code, whereby the brain uses information about the relative activity of hair cells across the whole
basilar membrane to help determine the pitch you hear. Second, the hair cell hairs move in time with the incoming sound wave, so auditory nerve axons fire synchronously
with the sound-wave peaks, which happen regularly at the sound's repetition rate. This provides a temporal code, whereby the brain uses the timing of the action
potentials in the auditory nerve to help determine the pitch you hear.
• Timbre depends, in part, on the relative amounts of different frequency components in a sound and this, like pitch, depends on the relative activity of hair cells across the
whole basilar membrane.
• How we determine the location of a sound has also been studied extensively. As in vision, there are monaural (one ear) and binaural (two ear) cues to location. Pinnas
intricate folds alter sound, emphasizing some frequency components over others, depending on where the sound is coming from. You have learned to interpret these
changes as indicating a sound's location. Second, the speed of sound is much slower than the speed of light. Sounds arrive a little sooner at the ear nearer to the source
than at the far ear. This time difference is effective for indicating the location of lower-frequency components of a sound, even when it is only a little off to one side. Third,
the higher-frequency components of a sound are more intense in the ear close to the sound than in the ear farther away, because the listener's head blocks higher
frequencies. The farther a sound is off to the side, the greater the between-ear difference in the level of these high-frequency components.

Sometimes you may find yourself turning your head from side to side to localize a sound. By doing this, you are changing the relative intensity and timing of sound waves arriving
in your ears and collecting better information about the likely source of the sound. Turning your head also allows you to use your eyes to locate the source of the sound — and,
as you learned earlier in this chapter, your visual system is much better at pinpointing the location of things than your auditory system is.

In hearing, your auditory system first analyzes a complex sound wave into its component frequencies, which activate different regions of the basilar membrane and auditory
nerve. Then the brain has to figure out which frequency components belong together in a single source (perceptual grouping) and which frequency components belong to
different sources (perceptual segregation). Just as in vision, our perceptual system organizes and automatically delivers to us the interpretation that is the simplest and most
meaningful, consistent with expectations based on experience.

The Gestalt rules you learned about earlier in this chapter also apply to sound. For example, sounds that are similar in the physical properties of frequency or intensity, or that
are similar in the perceptual attributes of loudness, pitch, timbre, or location, are grouped together into one source, as are sounds that occur close together in time (proximity).
Furthermore, sounds that start together and stop together (like the different frequencies emitted simultaneously by a musical instrument, or a voice) are perceived as coming
from the same source.

Hearing loss has two main causes. Conductive hearing loss arises because the eardrum or ossicles are damaged to the point that they cannot conduct sound waves effectively to
the cochlea. The cochlea itself, however, is normal, making this a kind of “mechanical problem” with the moving parts of the ear: the hammer, anvil, stirrup, or eardrum. In many
cases, medication or surgery can correct the problem. Sound amplification from a hearing aid also can improve hearing through conduction via the bones around the ear directly
to the cochlea.

Sensorineural hearing loss is caused by damage to the cochlea, the hair cells, or the auditory nerve, and it happens to almost all of us as we age. It has two main effects:
Sensitivity decreases so sounds have to be more intense to be heard, and acuity decreases so sounds smear together on the basilar membrane, making voices harder to
understand, especially if other sounds are present. Sensorineural hearing loss has many causes, including genetic disorders, premature birth, infections, medications,
accumulated damage from sound exposure (particularly intense sounds), and aging (these last two causes are hard to tease apart since older people have been exposed to sound
for longer). Hearing aids amplify sounds, helping with decreased sensitivity, but, unlike glasses, which can fix blurry vision, hearing aids cannot fix the acuity problem.

When hearing loss is severe, a cochlear implant may restore some hearing. A cochlear implant is an electronic device that replaces the function of the hair cells. The external
parts of the device include a microphone and a processor and transmitter (a little bigger than a two-dollar coin), worn on the scalp near the ear. The implanted parts include a
receiver just inside the skull and a thin wire containing electrodes inserted into the cochlea to stimulate the auditory nerve. Sound picked up by the microphone is transformed
into electric signals by the processor, which is essentially a small computer. The signal is transmitted to the implanted receiver, which activates the electrodes in the cochlea.
Cochlear implants are now in routine use and can improve hearing to the point that the wearer can understand speech, although background sound still poses a real challenge.

2. The Body Senses

Vision and hearing provide information about the world at a distance. By responding to light and sound energy in the environment, these “distance” senses allow us to identify
and locate the objects and people around us. In comparison, the body senses, also called somatosenses (soma from the Greek for “body”), are up close and personal. Haptic
perception is the active exploration of the environment by touching and grasping objects with our hands. We use sensory receptors in our muscles, tendons, and joints as well as
a variety of receptors in our skin to get a feel for the world around us.

Touch begins with the transduction of skin sensations into neural signals. Receptors located under the skin’s surface enable us to sense pain, pressure, texture, pattern, or
vibration against the skin. Each receptor has a tactile receptive field, a small patch of skin that relates information about pain, pressure, texture, pattern, or vibration to a
receptor. These specialized cells work together to provide a rich tactile (from Latin, “to touch”) experience when you explore an object by feeling it or attempting to grasp it. In
addition, thermoreceptors, nerve fibres that sense cold and warmth, respond when your skin temperature changes. All these sensations blend seamlessly together in perception,
of course, but detailed physiological studies have successfully isolated the parts of the touch system.

There are three important principles regarding the neural representation of the body’s surface. First, the left half of the body is represented in the right half of the brain and vice
versa. Second, different locations on the body send sensory signals to different locations in the somatosensory cortex in the parietal lobe. Just as more of the visual brain is
devoted to foveal vision, where acuity is greatest, more of the tactile brain is devoted to parts of the skin surface where sensitivity to fine spatial detail (acuity) is greatest.
Regions such as the fingertips and lips have high acuity, whereas acuity is lower in areas such as the calf. You can test this yourself: If you put two chopsticks together so that
their tips are about a centimetre apart, and gently press them into the skin of your fingertip, you should be able to tell that there are two tips, not one. If you do the same on
your calf, how far apart do the tips have to be before you can tell them apart? Third, there is mounting evidence for a distinction between “what” and “where” pathways in
touch, analogous to similar distinctions we’ve already considered for vision and hearing. The “what” system for touch provides information about the properties of surfaces and
objects; the “where” system provides information about the location in external space that is being touched or a location on the body that is being stimulated.

Does the possibility of a life free from pain seem appealing? Although pain is arguably the least pleasant of sensations, this aspect of touch is among the most important for
survival: Pain indicates damage or potential damage to the body. Without the ability to feel pain, we might ignore infections, broken bones, or serious burns. Congenital
insensitivity to pain, a rare inherited disorder that specifically impairs pain perception, is more of a curse than a blessing: Children who experience this disorder often accidentally
mutilate themselves (e.g., biting into their tongues or gouging their skin while scratching) and are at increased risk of dying in childhood.

Tissue damage is transduced by the free nerve endings that sense painful stimuli. In addition, researchers have distinguished between fast-acting A-delta fibres, which are axons
that transmit initial sharp pain, and slower C fibres, which are axons that transmit the longer-lasting, duller persistent pain.

The pain-withdrawal reflex is coordinated by the spinal cord. No brainpower is required when you touch a hot stove; you retract your hand almost instantaneously. But neural
signals for pain — such as wrenching your elbow as you brace yourself to keep from falling — travel to two distinct areas in the brain and evoke two distinct psychological
experiences. One pain pathway sends signals to the somatosensory cortex, identifying where the pain is occurring and what sort of pain it is (sharp, burning, or dull). The second
pain pathway sends signals to the motivational and emotional centres of the brain, such as the hypothalamus and amygdala, as well as to the frontal lobe. This is the aspect of
pain that is unpleasant and motivates us to escape from, or relieve, the pain.

Interestingly, fMRI has demonstrated that some of the brain areas that respond to our own physical pain, particularly in the frontal lobe, also respond when we witness others
experiencing pain and feel concern for them. Precisely what common features of experiencing pain and witnessing pain result in this overlap, and whether or not the overlap
extends to perceiving the social pain (such as embarrassment or social exclusion) of others, are areas of active research.

Pain typically feels as though it comes from the site of the tissue damage that caused it. If you burn your finger, you will perceive the pain as originating there. But we have pain
receptors in many areas besides the skin, such as around bones and within muscles and internal organs as well. When pain originates internally, in a body organ, for instance, we
can feel it on the surface of the body. This kind of referred pain occurs when sensory information from internal and external areas converges on the same nerve cells in the spinal
cord. One common example is a heart attack: Victims often feel pain radiating from the left arm rather than from inside the chest.

Pain intensity cannot always be predicted solely from the extent of the injury that causes the pain. For instance, turf toe sounds like the mildest of ailments; it is pain at the base
of the big toe as a result of bending or pushing off repeatedly, as a runner or football player might do during a sporting event. This small-sounding injury in a small area of the
body can nonetheless sideline an athlete for a month with considerable pain. On the other hand, you’ve probably heard a story or two about someone treading bone-chilling
water for hours on end, or dragging their shattered legs a mile down a country road to seek help after a tractor accident, or performing some other incredible feat despite
searing pain and extensive tissue damage. Pain type and pain intensity show a less-than-perfect correlation, a fact that intrigues researchers.

One influential account of pain perception is known as the gate-control theory, which holds that signals arriving from pain receptors in the body can be stopped, or gated, by
interneurons in the spinal cord via feedback from the skin or from the brain. Pain can be gated by the skin receptors, for example, by rubbing the affected area. Rubbing your
stubbed toe activates neurons that “close the gate” to stop pain signals from travelling to the brain. Pain can also be gated from the brain, which can modulate the activity of
pain-transmission neurons.

The brain’s feedback to the spinal cord comes from a region in the midbrain called the periaqueductal grey (PAG). Under extreme conditions, such as high stress, naturally
occurring endorphins can activate the PAG to send inhibitory signals to neurons in the spinal cord that then suppress pain signals to the brain, thereby modulating the experience
of pain. The PAG also responds to the action of opiate drugs, such as morphine.

A different kind of feedback signal from the brain can increase the sensation of pain. This system is activated by events such as infection and learned danger signals. When we are
quite ill, what we otherwise might experience as mild discomfort can feel quite painful. This pain facilitation signal presumably evolved to motivate people who are ill to rest and
avoid strenuous activity, allowing their energy to be devoted to healing.

Although some details of the gate-control theory of pain have been challenged, a key concept underlying the theory — that perception is a two-way street — has broad
implications. The senses feed information such as pain sensations to the brain, which processes these sensory data into perceptions, as you have learned in this chapter.* But, as
you have also learned, perceptions are not just shaped by sense data — they are affected by your knowledge, by your expectations, and by other factors such as your mood and
motivational state. Visual illusions and the Gestalt principles of closure (filling in what isn’t really there) provide some examples. This kind of top-down control also explains how
the brain influences the experience of touch and pain.

Shut your eyes and notice the position of your legs and feet, and arms and hands. Can you feel where they are in space? This is a sense that isn’t often talked about —
proprioception is your sense of bodily position. After all, you need some way to sense your position in physical space other than moving your eyes constantly to see the location
of your limbs. Your perception of the position (and movement) of your torso, limbs, hands, and feet in space depends on stimulation of receptors in the muscles, tendons, and
joints of your body, whereas information about which way is up and about head movement (to serve your balance) originates in the inner ear.

Sensory receptors provide the information we need to perceive the position and movement of our limbs, head, and body. These receptors also provide feedback about whether
we are performing a desired movement correctly and how resistance from held objects may be influencing the movement. For example, when you swing a baseball bat, the
weight of the bat affects how your muscles move your arm, as well as the sensation when the bat hits the ball. You can use muscle, joint, and tendon feedback about how your
arms actually moved to improve performance through learning.

Maintaining balance depends primarily on the vestibular system, the three fluid-filled semicircular canals and adjacent organs located next to the cochlea in each inner ear. The
semicircular canals are arranged in three perpendicular orientations and studded with hair cells that detect movement of the fluid when the head moves or accelerates. The
bending of the hairs of these cells generates activity in the vestibular nerve that is then conveyed to the brain. This detected motion enables us to maintain our balance.

Vision also helps us keep our balance. If you see that you are swaying relative to a vertical orientation, such as the walls of a room, you adjust your posture to keep from falling
over. Psychologists have experimented with this visual aspect of balance by placing people in rooms where the floor is stationary but the walls sway forwards and backwards. If
the room sways enough, people — particularly small children — will topple over as they try to compensate for what their visual system is telling them. When a mismatch
between the information provided by visual cues and vestibular feedback occurs, motion sickness can result.

3. The Chemical Senses

Olfaction is the least understood sense, and one of the most fascinating. Recall from the Neuroscience and Behaviour chapter that whereas the other senses connect first to the
thalamus, olfactory information enters the frontal lobe, amygdala, hippocampus, and other forebrain structures almost directly. This anatomy indicates that smell has a close
relationship with areas involved in emotional and social behaviour, as well as memory. Smell can signal whether a creature is unfriendly or friendly (or is a potential mate); it can
tell us whether a substance is probably delicious or is more likely to be toxic and dangerous; and it has an uncanny ability to remind us of long-ago places and people. Helen
Keller (1880–1968), a blind and deaf writer, said: “Smell is a potent wizard that transports you across thousands of miles and all the years you have lived. The odors of fruits waft
me to my southern home, to my childhood frolics in the peach orchard. Other odors, instantaneous and fleeting, cause my heart to dilate joyously or contract with remembered
grief”.

Countless substances release odours into the air. Just as natural lights and sounds are mixtures of wavelengths and frequencies, most natural odours (such as baking bread,
coffee, and farts) are actually mixtures of different odourant molecules. Odourants are chemicals such as hydrogen sulfide (which on its own smells like rotten eggs),
benzaldehyde (which smells like almonds), and vanillin (which gives vanilla its distinctive smell). Odourant molecules make their way into our noses, drifting in on the air we
breathe. Situated along the top of the nasal cavity is a mucous membrane called the olfactory epithelium, which contains about 10 million olfactory receptor neurons (ORNs),
receptor cells that transduce odourant molecules into neural impulses. Each ORN has receptors that bind to some odourants but not to others, as if the receptor is a lock and the
odourant is the key. Groups of ORNs sensitive to the same odourant send their axons from the olfactory epithelium into the olfactory bulb, a brain structure located above the
nasal cavity beneath the frontal lobes. Humans possess about 350 different ORN types that permit us to discriminate up to one trillion (!) odours through the unique patterns of
neural activity each odour evokes. This setup is similar to our ability to see a vast range of colours through only a small number of retinal receptor cell types or to feel a range of
skin sensations through only a handful of touch-receptor cell types.

Dogs and mice have up to 900 different ORN types, and up to 100 times as many ORNs; they have a correspondingly sharpened olfactory sensitivity (ability to detect odours) and
olfactory acuity (ability to discriminate among odours). Nevertheless, humans are sensitive to the smells of some substances in extremely small concentrations. For example, we
can sense mercaptan, a chemical compound that is added to natural gas to help detect gas leaks, at a concentration of just 0.0003 part per million. By contrast, acetone (nail
polish remover), something most people regard as pungent, can be detected only if its concentration is 15 parts per million or greater.

Just as the output of the final stage of retinal processing is retinal ganglion cell axons that form the optic nerve, the output of the final stage of olfactory bulb processing is in the
axons that form the olfactory nerve. The olfactory bulb sends outputs to various centres in the brain, including the parts that are responsible for controlling basic drives,
emotions, and memories. The relationship between smell and emotion explains why smells can have immediate, strongly positive or negative effects on us. Fortunately, sensory
adaptation is at work when it comes to smell, just as it is with the other senses. Whether the associations are good or bad, after just a few minutes the smell fades. Smell
adaptation makes sense: It allows us to detect new odours that may require us to act, but after that initial evaluation has occurred, it may be best to reduce our sensitivity to
allow us to detect other smells.

Our experience of smell is determined not only by bottom-up influences, such as odourant molecules binding to sites on ORNs, but also by top-down influences, such as our
previous experiences with an odour. Consistent with this idea, people rate the identical odour as more pleasant when it is paired with an appealing verbal label such as cheddar
cheese rather than an unappealing one such as body odour.

Smell may also play a role in social behaviour. Other animals can detect odours from pheromones, biochemical odourants emitted by other members of an animal’s species that
can affect its behaviour or physiology. Pheromones play an important role in reproductive and social behaviour in insects and in several mammalian species, including mice, dogs,
and primates. What about humans?

Since humans are mammals, it is possible — perhaps even probable — that we, like other mammals, both secrete and sense pheromones. However, evidence for human
pheromones from well-conducted and replicable experiments is scant. Studies have tended to use small sample sizes (which raises the risk of a result that isn’t true) and they
have not been successfully replicated. In order to prove the existence of human pheromones, we would first want to observe a behavioural or physiological response that could
be attributed to an airborne chemical, then identify and synthesize the active molecule, and ultimately confirm that humans have a specialized cell receptor and transduction
mechanism for that chemical. Tristan Wyatt, a zoologist at Oxford points out, “We become smellier in puberty, apart from other changes in sexual development (the growth of
more visible secondary sexual characteristics such as hair in our armpits and groins, for example). The changes in the smells we give off before and after puberty might give us
clues as to what molecules might be signals”.

Some evidence for a human pheromone comes from breastfeeding mothers and babies. Nursing in mammals is known to depend on smell. Human mothers produce a substance
from the glands around their nipples. If this secretion, taken from a lactating mother, is put under the nose of a 3-day-old newborn (not their own child), the baby responds with
head and mouth movements consistent with nursing behaviour. This suggests that one of the compounds in the secreted substance might be a pheromone that encourages
nursing behaviour in babies; indeed, the degree to which first-time mothers secrete this substance seems to correlate with the rate of weight gain in the babies in the first days
after birth.

The sense of taste evolved from the sense of smell. Whereas smell analyzes the chemical composition of things outside the body, taste does the same for things taken into the
body. One of the primary responsibilities of the chemical sense of taste is identifying things that are bad for you — as in poisonous and lethal. Some aspects of taste perception
are genetic, such as an aversion to extreme bitterness (which may indicate poison), and some are learned, such as an aversion to a particular food that once caused nausea. In
either case, the direct contact between the tongue and possible foods allows us to anticipate whether something will be harmful or nutritious.

The taste system contains just five main types of taste receptors, corresponding to five primary taste sensations: salt, sour, bitter, sweet, and umami (savoury). The first four are
probably familiar to you, but umami may not be. The umami receptor was discovered by Japanese scientists who attributed its stimulation to the rich savoury tastes evoked by
foods containing a high concentration of protein, such as miso, meats, and cheeses like blue cheese or old cheddar.

The tongue is covered with thousands of small bumps, called papillae, which are easily visible to the naked eye. Within most of the papilla are hundreds of taste buds, the organs
of taste transduction. The mouth contains 5,000 to 10,000 taste buds fairly evenly distributed over the tongue, the roof of the mouth, and the upper throat. Each taste bud
contains 50 to 100 taste receptor cells, and receptors for the different tastes are also pretty evenly distributed — any taste can be detected by any part of the tongue. Taste
perception fades with age. This may help explain why young children seem to be “fussy eaters,” since they may have greater sensitivity to taste sensations.

Each taste bud contains several types of taste receptor cells whose tips, called microvilli, react with tastant molecules in food. Just as odours are typically composed of a mixture
of odourants, foods usually contain a mixture of tastants. Salt receptors are most strongly activated by sodium chloride (table salt). Sour receptors respond to acids, such as
vinegar or lime juice. Bitter, sweet, and umami receptors also respond to tastants that give rise to these tastes in food.

Research suggests that there may be a sixth basic taste, fat, that is elicited by fatty acids and is distinct from the five primary taste sensations. The researchers noted that the
sensation people experience in response to fatty acids is not necessarily consistent with expectations of “fattiness,” so they proposed a new word for this taste sensation,
oleogustus. (Oleosus is Latin for “fatty” or “oily,” and gustus signifies “taste.”)

Taste experiences vary widely across individuals. About 50% of people report a mildly bitter taste in caffeine, saccharine, certain green vegetables, and other substances,
whereas roughly 25% report no bitter taste. Members of the first group are called tasters; members of the second group are called nontasters. The remaining 25% of people are
supertasters, who report that such substances, especially dark green vegetables, are extremely bitter, to the point of being inedible. Because supertasters tend to avoid fruits and
vegetables that contain tastes they experience as extremely bitter, they may be at increased health risk for diseases such as colon cancer. On the other hand, because they also
tend to avoid fatty, creamy foods, they tend to be thinner and may have decreased risk of cardiovascular disease. There is evidence that genetic factors contribute to individual
differences in taste perception, but much remains to be learned about the specific genes that are involved.

People also vary dramatically in their tolerance of food textures and spicy food. Some people hate the texture of eggplant (“slimy!”) or of bubble tea (“frogspawn!). Some people
really love hot spicy food; others do not. In addition to the five (or six) tastes to which receptors in the mouth are sensitive, information about the temperature, spiciness, and
texture (mouthfeel) of food is transduced and relayed from the mouth via touch receptors and thermoreceptors such as those discussed earlier.

Of course, the variety of taste experiences greatly exceeds the five basic taste receptors discussed here. Any food molecules dissolved in saliva evoke specific, combined patterns
of activity in the five taste receptor types. Furthermore, taste isn’t quite the same thing as flavour. Taste is the contribution made by receptors in your mouth alone. Taste and
smell collaborate to produce the complex perception of flavour. Have you ever eaten a meal while suffering through a severe head cold that blocks your sense of smell? In that
case, food tastes bland, right? The loss of smell and taste is a jarring and upsetting experience, as many people have learned for themselves during the COVID-19 pandemic.
Food perception is multisensory, involving taste, smell, and texture. Do these different sensory channels combine in one area of the brain to produce the perception of flavour?
Or are they each processed separately, such that perception of flavour depends on binding information across systems, rather like visual perception depends on binding
information from dorsal and ventral streams? Neuroimaging, particularly fMRI, has been very helpful in letting us get “inside the heads” (literally!) of people while they’re eating,
to begin to answer this question. This work indicates that areas sensitive to odours from food include not only primary olfactory cortex (near the amygdala) but also primary
gustatory (taste) cortex (in the frontal lobe) and the mouth region of primary somatosensory cortex (in the parietal lobe). Indeed, primary olfactory and gustatory cortex are
interconnected, and gustatory cortex passes “taste” information directly to olfactory cortex.

Multisensory effects on the perception of flavour also include visual and auditory influences. Many studies have shown that the colour of food and drinks can alter ratings of how
they taste, and similar effects have been observed even for packaging and labels. For example, in a study on beer tasting Barnett and Spence (2016) reported that participants
gave higher ratings of citrus and fruity tastes to a beer that was poured from a bottle with a yellow-green label designed to highlight the citrus notes in the beer, versus the same
beer poured from a bottle with a brown label or no label. You’ve probably eaten many times in loud environments, such as noisy restaurants or an airline cabin at 30,000 feet. In
a study that was designed to simulate the kind of loud background noise that you typically experience in an airplane cabin, Yan and Dando (2015) found that participants who
rated a variety of taste solutions gave lower sweet intensity ratings and higher umami taste intensity ratings in the noisy condition as compared to a control condition. More
recent research using similar levels of loud noise has shown that it affects how people experience drinking a cup of coffee: Participants reported that coffee tasted less bitter and
had less aroma in noisy than in quiet conditions.

One last point about flavour: It is a very easy thing to learn about! In the Learning chapter, we discuss how a strong, lifelong aversion to a food (e.g., hummus) can develop after
only one nasty experience with a tainted bowl of it. This makes sense — if something is bad for you once, it probably will be again, and you learn quickly to stay well away.
Learned preferences in food are also important in determining flavour, and they depend dramatically on culture and experience. “It’s an acquired taste” is something you may
have heard about a food you find disgusting: Acquired tastes include toasted grasshoppers in Mexico (chapulines); fermented fish in Scandinavia (lutefisk, hakarl, surströmming);
fetal ducks still in the shell in Vietnam and Philippines (balut) — even cheese in many parts of the world. One person’s “yecch!” is another person’s “yummy!”