Notes Cognitive Neuroscience The Structure of Neurons - nervous system is composed of two main classes of cells: neurons and glial cells - - - - Neurons (100 billion / 10^11) - neurons are the basic signaling units that transmit information throughout the nervous system - neurons take in information, make a “decision” about it following some relatively simple rules, and then, by changes in their activity levels, pass it along to other neurons - vary in their form, location, and inter- connectivity within the nervous system (Figure 2.2), and these variations are closely related to their functions - standard cellular components found in almost all eukaryotic cells are found in neurons as well - neuron is composed of three main parts: a cell body, dendrites, and an axon § cell body contains the cellular machinery for the production of proteins and other cellular macromolecules § dendrites and axon are extensions of the cell membrane and contain cytoplasm continuous with that in the cell body the neuron contains a nucleus, endoplasmic reticulum, ribosomes, mitochondria, Golgi apparatus, and other intracellular organelles cell membrane encases the cell body (also called soma) o contains the metabolic machinery that maintains the neuron (nucleus, endoplasmic reticulum etc.) o these structures are suspended in cytoplasm § salty intracellular fluid that is made up of a combination of ions, predominantly ions of potassium, sodium, chloride, and calcium, as well as molecules such as proteins neuron sits in a bath of salty extracellular fluid, which is also made up of a mixture of the same types of ions possess unique cytological features and physiological properties that enable them to transmit and process information rapidly Dendrites o branching extensions of the neuron that receive inputs from other neurons o take many varied and complex forms, depending on the type and location of the neuron o have specialized processes called spines § little knobs attached by small necks to the surface of the dendrites § where the dendrites receive inputs from other neurons Axons o a single output structure that extends from the axon hillock at the base of the cell soma, away from the cell body o Electrical signals travel down the axon toward the axon terminals, where the neuron transmits the signal to other neurons or other cell types (via dendrites and other axons) Axon Hillock o Is a specialized part of the cell body that is the connection to the axon o The last site in the cell body where membrane potentials propagated from synaptic inputs are summated before being transmitted to the axon Sagittal section through a cerebellar cortex showing a Purkinje cell Neurons located in the ventral horn of the spinal cord send their axons out the ventral root to make synapses on muscle fibers Glial Cells (1 trillion / 10^12) nonneural cells that serve various functions in the nervous system include providing structural support and electrical insulation to neurons, and modulating neuronal activity roughly as many glial cells in the brain as there are neurons Located throughout the nervous system, they may ac- count for more than half of the brain’s volume provide structural support helping to form the blood–brain barrier and aiding in the speed of information transfer have a previously unrecognized role in modulating neural activity are also the myelin formers in the nervous system central nervous system has three main types of glial cells: astrocytes, microglial cells, and oligodendrocytes Astrocytes large glial cells with round or radially symmetrical forms surround neurons and are in close contact with the brain’s vasculature makes contact with blood vessels at specializations called end feet (permit the astrocyte to trans- port ions across the vascular wall) create a barrier, called the blood–brain barrier (BBB), between the tissues of the central nervous system and the blood restricts passing of most bacteria, large hydrophilic molecules in the blood in the neural tissue allows small hydrophobic molecules such as oxygen, carbon dioxide, and hormones to enter active role in brain function Microglial Cells small and irregularly shaped come into play when tissue is damaged devouring and removing damaged cells can proliferate even in adults Oligodendrocytes and Schwann cells in the central nervous system, oligodendrocytes form myelin in the peripheral nervous system, Schwann cells form myelin create myelin by wrapping their cell membranes around the axon in a concentric manner during development and maturation cytoplasm in that portion of the glial cell is squeezed out leaving primarily the lipid bilayer of the glial cell sheathing the membrane Key Points the Structure of Neurons: Neurons and glial cells make up the nervous system. Neurons are the cells that transmit information throughout the nervous system. Most neurons consist of a cell soma (body), axon, and dendrites. Neurons communicate with other neurons and cells at specialized structures called synapses, where chemical and electrical signals can be conveyed between neurons. An astrocyte is a type of glial cell that helps form the blood–brain barrier. Astrocytes have an active role in modulating neural activity. Glial cells aid in the speed of information transfer by forming myelin around the axons of the neurons. An oligodendrocyte is a type of glial cell that forms myelin in the central nervous system. A Schwann cell is a type of glial cell that forms myelin in the peripheral nervous system. As part of the immune response of the nervous system, microglial cells are phagocytic cells that engulf damaged cells. Myelin is a good electrical insulator because the layers of cell membrane are composed of lipid bilayers, which are themselves poor electrical conductors Neuronal Signaling - Neurons receive, evaluate, and transmit information - Two types of transportation - Information is transferred across synapses from one neuron to the next, or from a neuron to a non-neuronal cell such as those in muscles or glands - being received at synapses on dendrites, conducted with- in the neuron, transmitted down the axon, and passed along at synapses on the axon terminals - Within a neuron, transferring information involves changes in the electrical state of the neuron as electrical currents flow through the volume of the neuron - Between neurons, information transfer occurs at synapses, typically mediated by chemical signaling molecules (neurotransmitters) but, in some cases, also by electrical signals - Regarding information flow, neurons are referred to as either presynaptic or postsynaptic in relation to any particular synapse - Most neurons are both presynaptic and postsynaptic Presynaptic - A neuron is presynaptic when their axon makes a connection onto other neurons Postsynaptic A neuron is postsynaptic when other neurons make a connection onto their dendrites The Membrane Potential process of signaling has several stages energy is needed to generate the signals this energy is in the form of an electrical potential across the neuronal membrane o electrical potential is the voltage inside the neuron versus out- side the neuron these two voltages depend on the concentrations of potassium, sodium, and chloride ions as well as on charged protein molecules both inside and outside of the cell voltage difference across the neuronal membrane in the resting state is typically −70 millivolts (mV) inside (known as resting membrane potential) o means that the neuron has at its disposal a kind of battery o stored energy can be used to do work lipid membrane maintains the separation of intracellular and extracellular ions and electrical charge that ultimately permits neuronal communication membrane is peppered with transmembrane proteins that serve as conduits for ions to move across the neuronal membrane two main types of these proteins: ion channels and ion pumps Ion Channels proteins with a pore through their centers they allow certain ions to flow down their concentration gradients transmembrane passageways created by ion channels are formed from the three-dimensional structure of these proteins o hydrophilic channels selectively permit one type of ion to pass through the membrane the ion channels that are found in neurons contain … o … either sodium, potassium, calcium, or chloride ions (Na+, K+, Ca2+, and Cl−) (figure 2.8) - - permeability: extent to which a particular ion can cross the membrane through a given ion channel membrane permeability to K+ is larger neurons can change the permeability of their membranes gated ion channels: ion channels that are capable of changing their permeability for a particular ion o open or close based on changes in nearby transmembrane voltage o open or close as a response to chemical or physical stimuli non-gated ion channels: are unregulated, and hence always allow the associated ion to pass through Ion pumps use energy to actively transport ions across the membrane against their concentration gradients o from regions of low concentration to regions of higher concentration there are con- centration gradients of different ions across the neuronal membrane Na+ and Cl− concentrations are greater outside of the cell - K+ concentrations are greater inside the cell - neurons use a Na+/K+ pump that pumps Na+ ions out of the cell and K+ ions into the cell - this process requires energy - Each pump is an enzyme that hydrolyzes adenosine triphosphate (ATP) - For each molecule of ATP that is hydrolyzed, the resulting energy is used to move three Na+ ions out of the cell and two K+ ions into the cell (figure 2.8 and 2.9) - concentration gradients create forces of the unequal distribution of ions - force of the Na + concentration gradient wants to push Na + from an area of high concentration to one of low concentration (outside to inside) - K+ con- centration gradient acts to push K+ from an area of high concentration to an area of low concentration (inside to outside) - inside and outside voltages are different because the membrane is more permeable to K+ than to Na + - electrical gradient: force of the K+ concentration gradient pushes some K+ out of the cell, leaving the inside of the neuron slightly more negative than the outside o o each K+ ion carries one unit of positive charge out of the neuron as it moves across the membrane These two gradients (electrical and ionic con- centration) are in opposition to one another with respect to K+ (figure 2.10) The Action Potential electrotonic conduction o Passive current conduction o maximum distance a passive current will flow is only about 1 millimeter o diminishes with distance from its origin—the synapse, in this case action potential o a rapid depolarization and repolarization of a small region of the membrane caused by the opening and closing of ion channels o doesn’t decrement after only 1 millimeter o can travel for meters with no loss in signal strength, because they continuously regenerate the signal o giraffes and blue-whales o metabolically expensive, and it contributes to the inordinate amount of the body’s energy used by the brain voltage-gated ion channels o action potential is able to regenerate itself due to the presence o located in the neuronal membrane o are found at the spike-triggering zone in the axon hillock and along the axon spike-triggering zone o initiates the action potential o passive electrical currents that are generated sum together at the axon hillock o flows across the neuronal membrane in the spike-triggering zone, depolarizing the membrane o depolarization is strong enough (-70mV to -55mV) – then an action potential is triggered equilibrium potential o particular voltage at which there is no net flux of ions o result = the membrane is temporarily hyperpolarized, meaning that the membrane potential is even farther from the threshold required for triggering an action potential hyperpolarization o causes the K+ channels to close o o - - resulting in the membrane potential gradually returning to its resting state during this state -> voltage-gated Na+ channels are unable to open, and another action potential cannot be generated refractory period o lasts only a couple of milliseconds and has two consequences § the neuron’s speed for generating action potentials is limited to about 200 action potentials per second § the passive current that flows from the action potential can- not reopen the ion-gated channels that generated it saltatory conduction o appearance that the action potential is jumping down the axon at great speed, from one node of Ranvier to the next Key Points Neuronal Signaling: - The presynaptic cell is located before the synapse with respect to information flow; the postsynaptic cell is located after the synapse with respect to information flow. Nearly all neurons are both pre- and postsynaptic, since they both receive and transmit information. - The resting membrane potential is the difference in the voltage across the neuronal membrane during rest (i.e., not during any phase of the action potential). - The electrical gradient results from the asymmetrical distribution of ions across the membrane. The electrical difference across the membrane is the basis of the resting potential. - Ion channels are formed by transmembrane proteins that create passageways through which ions can flow. - Action potentials are an all-or-none phenomenon: The amplitude of the action potential does not depend on the size of the triggering depolarization, as long as that depolarization reaches threshold for initiating the action potential. - Ion channels can be either passive (always open) or gated (open only in the presence of electrical, chemical, or physical stimuli). Passive current conduction is called electrotonic conduction or decremental conduction. A depolarizing current makes the inside of the cell more positive and therefore more likely to generate an action potential; a hyperpolarizing current makes the inside of the cell less positive and therefore less likely to generate an action potential. Voltage-gated channels are of prime importance in generating an action potential because they open and close according to the membrane potential. Myelin allows for the rapid transmission of action potentials down an axon. Nodes of Ranvier are the spaces between sheaths of myelin where voltage-gated Na + and K+ channels are located and action potentials occur. Synaptic transmission - the transfer of a signal from the axon terminal to the next cell two major kinds of synapses— chemical and electrical—each using very different mechanisms for synaptic transmission Chemical Transmitters neurons send a signal to the cell across the synapse by releasing neurotransmitters into the synaptic cleft arrival of the action potential at the axon terminal leads to the depolarization of the terminal membrane o causing voltage-gated Ca2+ channels to open then triggers small vesicles containing neurotransmitter to fuse with the membrane at the synapse and release the transmitter into the synaptic cleft (1) transmitter diffuses across the cleft (2) binds with specific receptors embedded in the postsynaptic membrane (figure 2.12) (3) neurotransmitter binding induces a change in the receptor (4) opens specific ion channels and results in an influx of ions leading to either depolarization (excitation) or hyperpolarization (inhibition) of the postsynaptic cell (5) hyperpolarization of the postsynaptic neuron produces an inhibitory postsynaptic potential (IPSP) Synaptic Vesicles – stores various neurotransmitters that are release at the synapse. The release is regulated by a voltagedependent calcium channel o Vesicles – essential for propagating nerve impulses between neurons and are constantly recreated by the cell Synaptic Cleft – the gap between neurons and synapses Receptor – in the cell’s (neuron’s) membrane, receiving chemical input such as from neurotransmitters o Ionotropic – ion-channel receptors that accept the flow of neurotransmitters o Metabotropic – indirectly allow the flow of neurotransmitters § - - - Neurotransmitters (amino acids, biogenic amines, neuropeptides) Small-molecule: single amino acids o Acetylcholine (muscle - the chemical that motor neurons of the nervous system release to activate muscles o Monoamines – involved in the regulation of cognitive processes such as emotion, arousal and certain types of memory o Dopamine (pleasure and addiction) – the reward chemical. The release of dopamine “teaches” the brain that an activity or stimulus is beneficial. Overstimulation via dopamine can lead to addiction. o Serotonin (sleep, eating, depression) – mediates gut movements and the perceptions of resource availability § Amino acid – a simple organic compound containing both a carboxyl (COOH) and an amino (NH2) group § Glutamate – primarily used in learning and memory. The most abundant neurotransmitter in the Nervous System, accounting for 90% of all synaptic transmissions • Main brain excitatory – glutamate stimulates the brain § GABA – accounts for 90% of all inhibitions • Main brain inhibitory – GABA, Serotonin, and other inhibitory neurotransmitters do not stimulate the brain. Provides balance to the excitatory neurotransmitters o o Neuropeptides – large neurotransmitters, made up of strings of amino acids. Divided into 5 groups § Tachykinins – pain § Neurohypophyseal hormones – love, production of urine § Hypothalamic releasing hormones – stress, growth inhibition § Opioid peptides – endorphins, feel-good chemicals Gaseous – believed to be used to stimulate the production of other “messenger chemicals” Electrical Transmitters Neuronal membranes are touching at specializations called gap junctions The cytoplasms of the two neurons are essentially continuous These gap junction channels create pores connecting the cytoplasm of the two neurons (figure 2.14) Result = electrical changes in one are reflected instantaneously in the other Useful when information must be conducted rapidly, such as in the escape reflex of some invertebrates Groups of neurons with these synapses can activate muscles quickly Key Points Synaptic Transmission: - Synapses 10^15/per neuron are the locations where one neuron can transfer information to another neuron or specialized non-neuronal cell. They are found on dendrites and at axon terminals but can also be found on the neuronal cell body. - Chemical transmission results in the release of neurotransmitters from the presynaptic neuron and the binding of those neurotransmitters on the postsynaptic neuron, which in turn causes excitatory or inhibitory postsynaptic potentials (EPSPs or IPSPs), depending on the properties of the postsynaptic receptor. - Classes of neurotransmitters include amino acids, biogenic amines, and neuropeptides. - Neurotransmitters must be removed from the receptor after binding. This removal can be accomplished by (a) active reuptake back into the presynaptic terminal, (b) enzymatic breakdown of the transmitter in the synaptic cleft, or (c) diffusion of the neurotransmitter away from the region of the synapse. - Electrical synapses are different than chemical synapses as they operate by passing current directly from one neuron (presynaptic) to another neuron (postsynaptic) via specialized channels in gap junctions that connect the cytoplasm of one cell directly to the other. Overview of the Nervous System Structure Nervous System Structure - composed of the central nervous system (CNS), brain and spinal cord, and the peripheral nervous system (PNS), nerves (bundles of axons and glia) and ganglia (clumps of nerve cell bodies) outside of the CNS Peripheral nervous system (PNS): - courier network delivers sensory information to CNS and carries motor commands from CNS to muscles - Consisting of the nerves (bundles of axons and glia) and ganglia (clumps of nerve cell bodies) outside of the CNS - Activities of PNS are accomplished through 2 systems: - somatic motor system - controls voluntary muscles - - autonomic motor system - controls visceral functions - involved in controlling the involuntary action of smooth muscles (the heart and various glands) Autonomic nervous system has 2 subdivisions: sympathetic and parasympathetic branches (operate antagonistically e.g. sympathetic increases heart rate, parasympathetic slows it) sympathetic system o uses the neurotransmitter norepinephrine § activation of this system increases heart rate, diverts blood from the digestive tract to the somatic musculature and prepares the body for action • flight • fight parasympathetic system o uses acetylcholine as its transmitter o activation of this system slows heart rate, stimulates digestion and in general helps the body with functions germane to maintaining the body Central nervous system (CNS) made up of the delicate brain and spinal cord o Each encased in its protective, bony shell and suspended in a sea of cerebrospinal fluid (CSF) both the brain and the spinal cord are covered with three protective membranes – meninges outer membrane is the thick mater, the middle is the arachnoid mater and the inner and most delicate is the pia mater which firmly adheres to the surface of the brain in the CNS neurons are bunched together in various ways o Two of the most common organizational clusters are in a nucleus or in a layer § a nucleus is a relatively compact arrangement of nerve cell bodies and their connections, ranging from hundred to millions of neurons with similar inputs and outputs • Located throughout both the brain and the spinal cord o The outer layer of the brain – cerebral cortex § has billions of neurons § arranged in layers if thin sheets, folded across the surfaces of the cerebral hemisphere like a handkerchief § cortex has a thin grayish layer overlaying the whitish interior o The gray matter § Is composed of neuronal cell bodies § consisting of neuronal bodies § resembles a butterfly with two se § ventral horn contains the large motor neurons that project to muscles § dorsal horn contains sensory neurons and interneurons § interneurons project to motor neurons on the same (ipsilateral) and opposite (contralateral) sides of the spinal cord to aid in the coordination of limb movements § surrounds the central canal, which is an anatomical ex- tension of the ventricles in the brain and contains cerebrospinal fluid o The white matter § Consist of axons and glial cells § peripheral region is made up of white matter tracts § tracts of ascending sensory information and descending motor signals Key Points Overview of the Nervous System Structure: The central nervous system consists of the brain and spinal cord. The peripheral nervous system consists of all nerves and neurons outside of the central nervous system. The autonomic nervous system is involved in controlling the action of smooth muscles, the heart, and various glands. It includes the sympathetic and parasympathetic systems. The sympathetic system uses the neurotransmitter norepinephrine. This system increases heart rate, diverts blood from the digestive tract to the somatic musculature, and prepares the body for fight-or-flight responses by stimulating the adrenal glands. The parasympathetic system uses acetylcholine as a neurotransmitter. It is responsible for decreasing heart rate and stimulating digestion. Groups of neurons are called ganglia. The cerebral cortex is a continuous sheet of layered neurons in each hemisphere. The axons of cortical neurons and subcortical ganglia travel together in white matter tracts that interconnect neurons in different parts of the brain and spinal cord. The corpus callosum is the main fiber tract that connects the two hemispheres of the brain. The Spinal Cord (figure 2.19) - Takes in sensory information from the body’s peripheral sensory receptors, relays it to the brain, and conducts the final motor signals from the brain to the muscles - Each level of the spinal cord has a reflex pathway, such as the knee-jerk reflex mentioned earlier - Runs from the brainstem at about the first spinal vertebrae to its termination in the cauda equina (“horse tail”) - The cauda equina is enclosed in the bony vertebral column – a stack of separate bones (vertebrae) that extend from the base of the skull to the fused vertebrae at the coccyx (tailbone) - vertebral column is divided into sections: cervical, thoracic, lumbar, sacral, and coccygeal - spinal cord is similarly divided (excluding the coccygeal region, since we no longer have tails) into 31 segments - each segment has a right and a left spinal nerve that enters and exits from the vertebral column through openings called foramen - each spinal nerve has both sensory and motor axons: one afferent neuron carries sensory input through the dorsal root into the spinal cord, and the other efferent neuron carries motor output through the ventral root away from it A Guided Tour of the Brain when we see a brain, the cerebral cortex, the outer layer, is most prominent for the brain, the cerebral cortex is the frosting on the cake—it’s the last thing to develop from an evolutionary at the base of the brain, are structures that are found in most vertebrates and have evolved for hundreds of millions of years these parts of the brain control our most basic survival functions, such as breathing, heart rate, and temperature - - - - - the prefrontal cortex, which is found only in mammals, is evolutionarily the youngest part of our brain damage to the prefrontal cortex may not be immediately fatal, but it will likely affect such things as our ability to make decisions as well as other behaviors that we consider to be most advanced in humans the head is merely an appendage to the body, so the terms that are used to describe the orientation of the head and its brain are in relation to the body let’s first picture the body of an Australian shepherd, looking off to the left o the front end is the rostral end, meaning “nose.” o the opposite end is the caudal end, the “tail.” o along his back is the dorsal surface, just like the dorsal fin is on the back of a shark. o the bottom surface along the dog’s belly is the ventral surface. o we can refer to the dog’s nervous system by using the same coordinates o the part of the brain toward the front is the rostral end (toward the frontal lobes) o the posterior end is the caudal end (toward the occipital lobe) o along the top of his head is the dorsal surface, and the bottom surface of the brain is the ventral surface. we humans are atypical animals because we stand upright and, therefore, tilt our heads forward in order to be parallel with the ground - thus, the dorsal surface of the body and brain are now at right angles to each other - in humans, we also use the terms superior and inferior to refer to the top and bottom of the brain, respectively - when we consider the spinal cord, the coordinate systems align with the body axis - thus, in the spinal cord, rostral means “toward the brain,” just as it does in the dog - if we slice the brain from nose to tail, that is a sagittal section - when that slice is directly through the middle, it is a midsagittal or medial section - If it is off to the side, it is a lateral section - If sliced from top to bottom, separating the front of the brain from the back, we have made a coronal section If we slice in a plane that separates dorsal from ventral, that is known as either an axial, transverse, or horizontal section The chambers of the Mind neurons in the brain are functional units, and that how they are interconnected yields specific circuits for the support of particular behaviors early anatomists, believing that the head contained the seat of behavior, examined the brain to see where the conscious self (soul, if you wish) was located Some chambers in the brain seemed to be empty (except for some fluid) and thus were possible containers for higher functions - ventricles brain weighs a considerable amount but has little or no structural support; there is no skeletal system for the brain to overcome this potential difficulty, the brain is immersed in a fluid called cerebrospinal fluid (CSF) this fluid allows the brain to float to help offset the pressure that would be present if the brain were merely sitting on the base of the skull o also reduces shock to the brain and spinal cord during rapid accelerations or decelerations, such as when we fall or are struck on the head ventricles inside the brain are continuous with the CSF surrounding the brain - largest of these chambers are the lateral ventricles, which are connected to the third ventricle in the brain’s midline cerebral aqueduct joins the third to the fourth ventricle in the brainstem below the cerebellum CSF is produced in the lateral ventricles and in the third ventricle by the choroid plexus, an out- pouching of blood vessels from the ventricular wall (hence CFS is similar to blood) CSF is a clear fluid containing proteins, glucose, and ions, especially potassium, sodium, and chloride o slowly circulates from the lateral and third ventricles through the cerebral aqueduct to the fourth ventricle and on to the subarachnoid space surrounding the brain, to be reabsorbed by the arachnoid villi in the sagittal sinus (the large venous system located between the two hemispheres on the dorsal surface; not shown) The Brainstem: Medulla, Pons, Cerebellum and Midbrain three main parts: the medulla (myelencephalon), the pons and cerebellum (metencephalon), and the midbrain (mesencephalon) these three sections form the central nervous system between the spinal cord and the diencephalon though the brainstem is rather small compared to the vast bulk of the forebrain, it plays a starring role in the brain - brainstem contains groups of (1)motor nuclei, (2)sensory nuclei, (3)nuclei of widespread modulatory neurotransmitter systems, (4)white matter tracts of ascending sensory information, (5)descending motor signals - damage to the brainstem = life threatening - brainstem nuclei control respiration and global states of consciousness such as sleep and wakefulness - medulla, pons, and cerebellum make up the hindbrain Midbrain Brainstem Hindbrain Mesencephalon Metencephalon Myelencephalon Cerebellum Pons Medulla Medulla - brainstem’s most caudal portion is the medulla - continuous with the spinal cord (figure 2.21) - essential for life - houses the cell bodies of many of the 12 cranial nerves, providing sensory and motor innervations to the face, neck, abdomen, and throat (including taste) as well as the motor nuclei - - that innervate the heart controls vital functions such as respiration, heart rate, and arousal all of the ascending somatosensory information entering from the spinal cord passes through the medulla via two bilateral nuclear groups gracile nuclei cuneate nuclei these projection systems continue through the brainstem to synapse in the thalamus en route to the somatosensory cortex the medulla is a relay station for sensory and motor information between the body and brain it is the crossroads for most of the body’s motor fibers it controls several autonomic functions, including the essential reflexes that determine respiration, heart rate, blood pressure, and digestive and vomiting responses Pons - - pons is the main connection between the brain and the cerebellum Sitting anterior to the medulla made up of a vast system of fiber tracts interspersed with nuclei Many of the cranial nerves synapse in the pons (include the sensory and motor nuclei from the face and mouth and the visuomotor nuclei controlling some of the extraocular muscles) important for some eye movements as well as those of the face and mouth responsible for generating rapid eye movement (REM) sleep Cerebellum - The cerebellum clings to the brainstem at the level of the pons - it is home to most of the brain’s neurons - it is a continuous layer of tightly folded neural tissue (like an accordion) - forms the roof of the fourth ventricle and sits on the cerebellar peduncles, which are massive input and output fiber tracts of the cerebellum - several gross subdivisions (1)cerebellar cortex (2)four pairs of deep nuclei, (3)the internal white matter - resembles the forebrain’s cerebral hemispheres - inputs from vestibular projections involved in balance, as well as auditory and visual inputs, also project to the cerebellum from the brainstem - output from the cerebellum originates in the deep nuclei ascending outputs travel to the thalamus and then to the motor and premotor cortex. other outputs project to nuclei of the brainstem, where they impinge on descending projections to the spinal cord - critical for maintaining posture, walking, and performing coordinated movements - not directly control movements; instead, it integrates in- formation about the body, such as its size and speed, with motor commands - then modifies motor outflow to effect smooth, coordinated movements - if cerebellum = damaged - your movements will be uncoordinated and halting, and you may not be able to maintain balance Midbrain (Diencephalon and Telencephalon) lies superior to the pons and can be seen only in a medial view surrounds the cerebral aqueduct, which connects the third and fourth ventricles dorsal portion consists of the tectum ventral portion is the tegmentum (covering) contains some of the cranial nerve ganglia and two other important structures superior colliculi plays a role in perceiving objects in periphery (the outer limits or edge of an area or object) orienting our gaze directly toward them and bringing them into sharper view inferior colliculi used for locating and orienting toward auditory stimuli red nucleus involved in certain aspects of motor co- ordination helps a baby crawl or coordinates the swing of your arms as you walk Key Points the Brainstem: Medulla, Pons, Cerebellum and Midbrain: The spinal cord conducts the final motor signals to the muscles, and it relays sensory information from the body’s peripheral receptors to the brain. The brainstem’s neurons carry out many sensory and motor processes, including visuomotor, auditory, and vestibular functions as well as sensation and motor control of the face, mouth, throat, respiratory system, and heart. The brainstem houses fibers that pass from the cortex to the spinal cord and cerebellum, and sensory fibers that run from spinal levels to the thalamus and then to the cortex. Many neurochemical systems have nuclei in the brainstem that project widely to the cerebral cortex, limbic system, thalamus, and hypothalamus. The cerebellum integrates information about the body and motor commands and modifies motor outflow to effect smooth, coordinated movements. - The Diencephalon: Thalamus and Hypothalamus - After leaving the brainstem, we arrive at the diencephalon, which is made up of the thalamus and hypothalamus - These subcortical structures are composed of groups of nuclei with interconnections to widespread brain areas Thalamus - Almost in the center of the brain and perched on top of the brainstem at the rostral end - The larger of the two diencephalon structures - Divided into two parts Right hemisphere Left hemisphere Both straddle the third ventricle The two parts are connected by a bridge of gray mater called the massa intermedia (figure 2.23) - referred to as the “gateway to the cortex” because all of the sensory modalities make synaptic relays in the thalamus before continuing to the primary cortical sensory receiving areas receives inputs from the basal ganglia, cerebellum, neocortex, and medial temporal lobe sends projections back to these structures to create circuits involved in many different functions thalamus is divided into several nuclei that act as specific relays for incoming sensory information lateral geniculate nucleus receives information from the ganglion cells of the retina and sends axons to the primary visual cortex medial geniculate nucleus receives information from the inner ear, via other brainstem nuclei in the ascending auditory pathway, and sends axons to the primary auditory cortex ventral posterior (medial and lateral) nuclei projects somatosensory information to the primary somatosensory cortex pulvinar nucleus Located at the posterior pole of the thalamus involved in attention and in integrative functions involving multiple cortical areas - - Hypothalamus main link between the nervous system and the endocrine system main site for hormone production and control two bumps seen on the ventral surface of the brain (mammillary bodies) belong to the small collection of nuclei and fiber tracks receives inputs from the limbic system structures and other brain areas job - control circadian rhythms (light–dark cycles) with inputs from the mesencephalic reticular formation, amygdala, and the retina controls the functions necessary for maintaining the normal state of the body (homeostasis) sends out signals that drive behavior to alleviate such feelings as (1)thirst, (2)hunger,(3) fatigue, and it controls body temperature and circadian cycles Key Points the Diencephalon: Thalamus and Hypothalamus: The thalamus is the relay station for almost all sensory information. The hypothalamus is important for the autonomic nervous system and endocrine system. It controls functions necessary for the maintenance of homeostasis. It is also involved in control of the pituitary gland. The pituitary gland releases hormones into the bloodstream where they can circulate to influence other tissues and organs (e.g., gonads). The Telencephalon: Limbic System and Basal Ganglia includes the cerebral cortex, the limbic system, and the basal ganglia forms a clump of structures found deep within the cerebral hemispheres nestled over and around the diencephalon Limbic System made up of the cingulate gyrus, the hypothalamus, anterior thalamic nuclei, and the hippocampus cingulate gyrus = a band of cerebral cortex that extends above the corpus callosum in the anterior–posterior direction and spans both the frontal and parietal lobes hippocampus = an area located on the ventromedial aspect of the temporal lobe neither anatomically nor functionally organized to the degree that other systems are in the brain includes the amygdala, the orbitofrontal cortex and parts of the basal ganglia group of neurons anterior to the hippocampus Basal Ganglia collection of nuclei bilaterally located deep in the brain beneath the anterior portion of the lateral ventricles voluntary motor responses, decision making its components are amygdala, striatum receive inputs from sensory and motor areas - striatum receives extensive feedback projections from the thalamus the basal ganglia have many dopamine receptors plays a crucial role in motivation and learning big role in reward-based learning and goaloriented behavior Key Points the Telencephalon: Limbic System and Basal Ganglia: The limbic system includes subcortical and cortical structures that are interconnected and play a role in emotion. The basal ganglia are involved in a variety of crucial brain functions, including action selection, action gating, reward-based learning, motor preparation, timing, task switching, and more. Cerebral Cortex crowning glory of the cerebrum is its outermost tissue, the cerebral cortex - made up of large sheets of (mostly) layered neurons draped and folded over the two symmetrical hemispheres sits over the top of the core structures including parts of the limbic system and basal ganglia, and surrounds the structures of the diencephalon infoldings of the cortical sheet are called sulci and gyri (figure 2.28) - folds of the human cortex serve several functions they enable more cortical surface to be packed into the skull total surface area of the human cerebral cortex is about 2,200 to 2,400 cm2 - - - - - cortex contains the cell bodies of neurons, their dendrites, and some of their axons cortex includes axons and axon terminals of neurons projecting to the cortex from other brain regions, such as the subcortical thalamus also contains blood vessels because of the high density of cell bodies, it appears grayish (figure 2.29) Gyri the crowns of the folded tissue that one observes when viewing the surface Dentate Gyrus – simple cortical regions that is an integral portion of the larger functional brain system called the hippocampal formation having a highly folded cortex brings neurons into closer three- dimensional relationships to one another reducing axonal distance and hence neuronal conduction time between different areas the cortex brings some nearby regions closer together cortex ranges from 1.5 to 4.5 mm in thickness, but in most regions, it is approximately 3 mm thick Inferior temporal gyrus – placed below the middle temporal sulcus, connected behind with the inferior occipital gyrus, one of the higher levels of the visual processing associated with the representation of complex object features Middle temporal gyrus – located between superior temporal gyrus and inferior temporal gyrus, recognition of known faces Inferior frontal gyrus – speech production Sulci furrow of a convoluted brain surface Central Sulcus – divides frontal from parietal lobes Lateral sulcus (Sylvian fissure) - separates the temporal lobe from the frontal and parietal lobes Cortical Layers are sheets of neurons neatly stacked on top of each other neurons of each layer are typically similar within a layer, but different between layers Molecular layer - contains the flattened dendritic trees of Purkinje cells, and the huge array of parallel fibers, from the granular layer, that penetrate the Purkinje cell dendritic trees at right angles External granular - thin layer consisting of numerous small, densely packed neurons External pyramidal - most superficial layer of cerebral cortex Internal granular - this layer receives the afferent connections from the thalamus and from other cortical regions and sends connections to the other layers Internal pyramidal - consists predominantly of the medium- sized and large pyramidal cells. It is the source of the output or corticofugal fibers Multiform (fusiform) layers - the deepest layer of the cortex that directly overlies the subcortical white matter. Brodmann’s areas (figure 2.31) - identified approximately 52 regions of the cerebral cortex - categorized and numbered according to differences in cellular morphology and organization - the numbering has more to do with the order in which Brodmann sampled a region than with any meaningful relation between areas - region of the cerebral cortex in the human or other primate brain, defined by its cytoarchitecture, or histological structure and organization of cells Lobes and Basic Functions cerebral hemispheres have four main divisions (lobes) that are best seen in a lateral view: the frontal, parietal, temporal, and occipital lobes (figure 2.30) skull bones them- selves are named for their locations lobes can usually be distinguished from one another by prominent anatomical landmarks such as pronounced sulci central sulcus divides the frontal lobe from the parietal lobe - Sylvian (lateral) fissure separates the temporal lobe from the frontal and parietal lobes occipital lobe is demarcated from the parietal and temporal lobes by the parieto-occipital sulcus left and right cerebral hemispheres are separated by the interhemispheric fissure (figure 2.28b) insula is located between the temporal and frontal lobe an island of folded cortex hidden deep in the lateral sulcus divided into the larger anterior insula and smaller posterior insula Occipital lobe (figure 2.35) - The job of the occipital lobes is vision - receives visual information relayed from the lateral geniculate nucleus of the thalamus - In humans, the primary visual cortex is on the medial sur- face of the cerebral hemispheres, extending only slightly onto the posterior hemispheric pole - The cortex in this area has six layers and begins the cortical coding of visual features like luminance, spatial frequency, orientation, and motion - Visual information from the outside world is processed by multiple layers of cells in the retina transmitted via the optic nerve to the lateral geniculate nucleus of the thalamus from there to V1—a path- way often referred to as the retinogeniculostriate (primary visual pathway) Parietal lobe (figure 2.34) - - receives sensory information from the out- side world, sensory information from within the body, and information from memory, and integrates it result in all sorts of odd deficits relating to sensation and spatial location Stimulating certain regions of the parietal lobe causes people to have “out of body” experiences Sensory information about touch, pain, temperature sense, and limb proprioception (limb position) is received via receptor cells on the skin converted to neuronal impulses that are conducted to the spinal cord after that conducted to the somatosensory relays of the thalamus (figure 2.34) Frontal lobe (figure 2.33) - plays a major role in the planning and execution of movements - two main subdivisions: the prefrontal cortex and the motor cortex - motor cortex sits in front of the central sulcus, beginning in the depths of the sulcus and extending anteriorly - auditory cortex lies in the superior part of the temporal lobe in a region - has a tonotopic organization, meaning that the physical layout of the neurons is based on the frequency of sound Temporal lobe audition, speech, perception, memory, emotions (auditory, visual, and multi- modal processing areas) temporal lobe lies underneath the temporal bone passage of time can be observed first in the graying of hair Key Points Cerebral Cortex & Lobes and Basic Function - Gyri are the protruding areas seen on the surface of the cortex; sulci, or fissures, are the enfolded regions of cortex. - Brodmann divided the brain into distinct regions based on the underlying cytoarchitectonic. - The lobes of the brain include the frontal, parietal, temporal, and occipital lobes. - The frontal lobe is for planning, cognitive control, and execution of movements. The parietal lobe receives sensory input about touch, pain, temperature, and limb position, and it is involved in coding space and coordinating actions. - The temporal lobe contains auditory, visual, and multi- modal processing areas. The occipital lobe processes visual information. The limbic lobe (not really a lobe) is involved in emotional processing, learning, and memory. - Topography is the principle that the anatomical organization of the body is reflected in the cortical representation of the body, both in the sensory cortex and motor cortex. - Association cortices are those regions of cortex outside the sensory specific and motor cortical regions. Association cortex receives and integrates input from multiple sensory modalities. Brain Evolution Brain Size to body size ratio Homo Sapiens have the largest brain size to body size ratio - Porpoise (and dolphin) are very close Brain size increase in hominids During evolution, humans have experienced an exponential increase in brain size In a simple life form: every position and function of a nerve cell is determined by genes Bigger Brains A single byte (or 8 bits) can represent 4 DNA base pairs. In order to represent the entire diploid human genome in terms of bytes, we can perform the following calculations: 9 9 6×10 base pairs/diploid genome x 1 byte/4 base pairs = 1,5×10 bytes 9 1,5×10 bytes = 1.5 Gigabytes Number of Neurons 100 billion (10^11) Arborization of dendrites At the same time the synaptogenesis is occurring, neurons of the brain are increasing the size of their dendritic arborization, extending their axons, and undergoing myelination Number of synapses 10 000 per neuron (10^15 synapses) Radial Unit Hypothesis Hypothesis First cells divide symmetrically: they keep the same function (Symmetrical division) After this they divide asymmetrically: one cell gets a different function (ie. Neuron) (Asymmetrical division) The cortex is formed from an inside-out matter New neurons migrate radially, you get exponentially more neurons Radial Unit Hypothesis First cells divide symmetrically in the ventricular Zone (VZ): keep the same function (Symmetrical division) After this, they divide asymmetrically: one cell gets a different function (ie. Neuron) and the differentiated cell migrates to the Cortical Plate (Asymmetrical division) The cortex is formed from an inside-out manner From the ventricular zone (VZ) to the cortical plate (CP), deeper layers first Neuron Migration happens along the radial glial cells Radial glial cells form radial units from the ventricular zone through the intermediate zone to the cortical plate These maintain their topography (relative locations) Symmetrical division for the first five to six weeks of gestation, the cells in the subventricular zone divide in a symmetrical fashion the result is exponential growth in the number of precursor cells Asymmetrical division At the end of six weeks, when there is a stockpile of precursor cells, asymmetrical division begins After every cell division One of the two cells formed becomes a migratory cell destined to be part of another layer The other cell of the two cells remains in the subventricular zone It continues to divide asymmetrically in the subventricular zone Radial glia cells Stretch from the subventricular zone to the surface of the developing cortex Their work does not end with development Radial glia cells are transformed into astrocytes in the adult brain Helping to form part of the blood-brain barrier Neurons migrate along the radial glia cells that form a pathway for the neurons Radial migration Cortical neurons are born and how they migrate radially from the ventricular zone toward the surface of the developing cortex Neurons migrate along the radial glial cells Radial glial highway is organized in a straight line from the ventricular zone to the cortical surface Radial unit Hypothesis - The fact that by prolonging the division, we can exponentially increase the number of neurons Provides the simplest explanation for our enormous increase in brain size Explains why the cortex is organized in columns Each unit is not enlarged, instead the number of units increases (**) Cortical column A principle unit of organization that has functional consequences and a development history The cortical columns that arise from (**) these groupings have functional and anatomical consequences in the adult Inside-out formation of cortex In which each cohort of neurons migrates past its cortical plate predecessors to form a more superficial layer Ventricular zone The cortical neurons that form other parts of the brain arise from precursor cells in the ventricular zone Subventricular Zone The cortical neurons arise from the subventricular zone Intermediate Zone Located between the ventricular zone and the cortical plate The white matter in this area is where neurons (created in the ventricular zone) migrate through in order to reach the cortical plate This zone is only present during carcinogenesis Eventually transform into adult white matter Cortical Plate In humans and many other species, the fetal brain is well developed and shows cortical layers, neuronal connectivity and myelination The fetal brain is already extremely complex, but far from completely being developed The first migrating neurons approach the surface of the developing cortex – point known as cortical plate Marginal Zone Predecessor for layer one of the cortex Marginal zone + cortical zone = 6 layers that form the cortex Synaptogenesis Substantial amount of that growth comes from synaptogenesis The formation of synapses and the growth of dendritic trees Early – in the deeper cortical layers Later – in more superficial layers Synaptogenesis – followed by synapse elimination (also called pruning) Synapse elimination/pruning Synaptic pruning is a competitive process Allows for learning development of higher cognitive functions The elimination of some synaptic contacts between neurons during development, including postnatally Eliminating the interconnections between neurons that are redundant, unused or do not remain functional Myelination Differential timing in human development Myelination of axons extends even further into life Axons in different cortical areas myelinate at different times Sensory and motor areas first Frontal and parietal areas last Myelination of frontal cortex continues way into adulthood :) Until age 25-30 Use it now (in your 20s) or never get it Ontogeny = Phylogeny Ontogeny recapitulates phylogeny Delayed maturation of the frontal lobes is an example of ontogeny following phylogeny A late addition in evolution means late development Brain Plasticity The brain also displays enormous potential for plasticity after complete development !!! your brain constantly changes!!! The brain can recruit other areas that are/were used for something else (Areas from the somatosensory cortex that are not used anymore are recruited by other parts) Reason why blind people can hear and feel better Term that refers to the brains ability to change and adapt as a result of experience Learning Mostly connectivity: synaptic changes Reorganization The brain can repair or reorganize after an accident Phantom limbs the ability to feel sensations and even pain in a limb or limbs that no longer exist Non-painful sensation: perception of movement and perception of external sensations (touch, temperature, pressure, vibration, itch) Pain sensations: range from burning and shooting pains to feelings of tingling “pins and needles” Only occurs in amputees Basic Genetics Chromosomes My chromosomes make me an individual Each chromosome consists of genes Genes A gene encodes a functional element (protein) A gene consists of a code (triplets of nuclei acids) DNA DNA is deoxyribonucleic Acid Consist of four bases: cytosine (C), guanine (G), adenine (A), or thymine (T) Base pairs: C – G and A – T When we reproduce, our DNA is combined with that of our partner Genotype The genes Phenotype How they are expressed Recombination and cross-over Combine two individuals Mutation Randomly alter a bit (usually low) Genetic Algorithms Genotype strings and individuals A genetic algorithm has a large number of strings/genotypes Each genotype is also called an individual Populations A collection of individuals is called a population Fitness value/function Each string/genotype/individual is assigned a fitness value according to phenotype This value can be normalized to a range between 0 and 1 You can use anything you like to calculate your fitness value Reproduction Cross-over You have various forms of cross-over The simples form is single-point cross-over 00110110 x 11001101 = 11000110 x 00111101 It is also possible to have multiple cross-over points 00110110 x 11001101 = 00101110 x 11010101 Inversion we select a subset of genes and invert the entire string in the subset 0123456789 x 0165432789 Mutation There are various types of mutation, depending on your representation Binary mutation: 001100 = 001110 Selection strategies Elitist selection Order individuals by fitness Select top -10, -20, … fittest individuals Pro: very straightforward to implement, very fast Con: each generation loses a lot of genetic information, “genetic degeneration” Roulette wheel selection The wheel is rotated The weakest individual has smallest share of the roulette wheel The fittest individual has largest share of the roulette wheel Pro: better than elitist selection Con: comes very close to elitist selection if fitness values are very unequal Tournament selection Select random k individuals (k=tournament size) From this selection: select the best individual Pro: Very straightforward; does not have problems of roulette wheel selection Con: a large k boils to elitist selection Genetic Algorithms Pros and Cons Advantages Widely acceptable Easy to implement Easy to parallelize Disadvantages Same local minima problem as with back-prop, but genetic algorithms are better suited to escape local minima Lots of variation (e.g. Selection, parameters) Sometimes difficult to find right encoding Genetic Algorithms applications Scheduling problems Financing Complex mathematical problems Engineering: NASA, Intel Chemistry Compilers Exoskeleton Airforce Strategy Advisor Airbus Cognitive Control Allows us to use our perceptions, knowledge, and goals to bias the selection of action and thoughts from a multitude of possibilities Allow us to override automatic thoughts and behavior and step out of the realm of habitual responses Give us cognitive flexibility, letting us think and act in novel and creative ways By being able to suppress some thoughts and activate others, we can simulate plans and consider the consequences of those plans We can plan for the future and troubleshoot problems Essential for purposeful goal- oriented behavior and decision making „you might want to stop at the doughnut store when heading to work in the morning, cognitive control mechanisms can override that sugary urge, allowing you to stop by the café for a healthier breakfast “ - Anatomical Orientation picture: first, which includes the lateral pre- frontal cortex and frontal pole supports goal-oriented behavior involved with planning, simulating consequences, and initiating, inhibiting, and shifting behavior second control system, which includes the medial frontal cortex essential role in guiding and monitoring behavior works in tandem with the prefrontal cortex monitoring ongoing activity to modulate the degree of cognitive control needed to keep behavior in line with goals Sensation and Perception Visible light spectrum segment of electromagnetic spectrum that the human eye can view human eye can detect wavelengths from 380 to 700 nanometers Longer wavelengths (red) Shorter wavelengths (blue) Eye Iris - the colored part - the black circular opening in the iris let’s light in pupil size is adjusted to filter the amount of light smaller for bright light larger for low light Pupil Cornea - - Lens - Retina - - a clear dome over the iris protective outer layer serves as barrier against dirt, germs, and other things that can cause damage also filters out some of the sun’s ultraviolet light as light enters your eye, it gets refracted by the corneas curved edge helps determine how well you can focus on objects close-up and far away Three main layers Epithelium Stops outside matter from getting into your eye Also absorbs oxygen and nutrients from tears Stroma Middle (thickest) layer lies behind epithelium Made up mostly of water and protein that give it an elastic but solid form Endothelium Single layer of cells on the very back of the stroma Stroma absorbs excess liquid and the endothelium pulls it out Without this function the stroma would become waterlogged Focuses light rays onto the retina Lens is transparent Can be replaced if necessary Lens deteriorates as we age – result – needing reading glasses Nerve layer lining the back of the eye Retina senses light and creates electrical impulses These impulses are sent through the optic nerve to the brain Sees pictures upside down Optic nerve Bundle of more than a million nerve fibers carrying visual messages from the retina to the brain In order to see we must have light and our eyes must be connected to the brain The brain controls what you see, since it combines images Retina sees pictures upside down – brain turns images right side up Reversal of the images that we see – like mirror in a camera Inverted projection on retina Images of the world that surrounds us are projected upside down onto our retina Image reversal – allows us tremendous peripheral vision and ability to see objects larger Retina Without reversal – limited view of our world (similar to viewing the world through a drinking straw) Blind Spot Small portion of the visual field of each eye Corresponds to the position of the optic disk within the retina No photoreceptors (rods or cones) in optic disk Therefore, no image detection in this area Blind spot of the right eye is located to the right of the center of vision Blind spot of the left eye is located to the left of the center of vision With both eyes open – blind spots not perceived because the visual fields of the two eyes overlap Optic Disk Can be seen in the back of the eye with an ophthalmoscope Located on the nasal side of the macula lutea Oval shape 1.5 mm in diameter Entry point into the eye for major blood vessels that serve the retina Fovea - Small, central pit composed of closely packed cones in the eye Located in the center of the macula lutea of the retina Responsible for sharp central vision Necessary in humans for activities for which visual detail – primary importance Surrounded by parafovea belt and perifovea outer region Employed for accurate vision in the direction where it is pointed Compromises less than 1% of the retinal size Takes up over 50% of the visual cortex Sees only the central two degrees of the visual field The farther away from the fovea the fewer receptor Central region of the retina that is densely packed with cone cells and provides high resolution visual information Receptors Rods Cones - Responsible for vision at low light levels (scotopic vision) Usually located around the boundary of the retina About 120 million photoreceptors out of the total 125 million photoreceptors in the human eye Outer segment is cylindrical – contain rhodopsin pigment (made up of vitamin A) Do not give color vision Do not have any differentiation If lack of the pigment in the rods = night blindness Fewer in number and are of cone shape Located in the center of the retina 5 million photoreceptors out of 125 million Outer segment is conical of cones (contain iodopsin pigment) Give color vision If lack of pigment – cause color blindness Chromatic (three types of pigment): Short wavelength - The blue part of the spectrum Medium wavelength The greenish region Long wavelength The reddish region Ganglion cells Final output neurons of the vertebrate retina Collect information about the visual world from bipolar cells and amacrine cells Process visual information Begins as light entering the eye Ganglion cells transmit it to the brain via their axons We have only 2 million ganglion cells to telegraph information from the retina Many rods feed into a single ganglion cell Each ganglion cell is innervated by only a few cones Bipolar cell Part of the retina Exist between photoreceptors (rod cells and cone cells) and ganglion cells They act directly or indirectly To transmit signals from the photoreceptors to the ganglion cells Receive synaptic input from either rods or cones, or both Horizontal cell Laterally interconnecting neurons having cell bodies in the inner nuclear layer of the retina Help integrate and regulate the input from multiple photoreceptor cells Receive input from multiple photoreceptor cells use that input to integrate signaling from different populations of photoreceptor cells Adjust the signals that will be sent to bipolar cells Amacrine cells Lie in the inner retina Make connections with bipolar cells and ganglion cells Create functional subunits within the receptive fields of many ganglion cells contribute to vertical communication within retinal layers Figure 5.23 shows how visual information is conveyed from the eyes to the central nervous system. Before entering the brain, each optic nerve splits into two parts. The temporal (lateral) branch continues to traverse along the ipsilateral side. The nasal (medial) branch crosses over to project to the contralateral side; this crossover place is called the optic chiasm. Lateral inhibition - The phenomenon in which a neurons response to a stimulus is inhibited by the excitation of a neighboring neuron - Observed in the retina and the lateral geniculate nucleus (lgn) of organisms - Makes neurons more sensitive to spatially varying of stimulus than to spatially uniform stimulus - Helps refine some somatosensory information - Only the neurons that are most stimulated and least inhibited respond - Plays an important role in visual perception by increasing the contrast and resolution of visual stimuli Contrast enhancement Defined as the manipulation and redistributing the image pixels in a linear or non-linear fashion Improve the separation of obscured structural variations in pixel intensity into a more visually differentiable structural distribution Required to increase the quality of low contrast images by expanding the dynamic range of input gray level contrast enhancement without disturbing other parameters of the image is one of the difficult tasks in image processing Visual system Comprises the sensory organ (eye) and parts of the central nervous system (the retina containing photoreceptor cells, the optic nerve, the optic tract and the visual cortex) Gives organisms the sense of sight Enabling the formation of several non-image photo response functions - Detects and interprets information from the optical spectrum perceptible to that species to “build a representation” of the surrounding environment Carries out number of complex tasks Reception of light and the formation of monocular neural representation Color vision The neural mechanisms underlying stereopsis and assessment of distances and between objects Identification of particular object of interest Motion perception Analysis and integration of visual information Pattern recognition Accurate motor coordination under visual guidance Eye - - Light entering the eye is refracted As it passes through the cornea Then passes through the pupil (controlled by the iris) Further refracted by the lens Cornea and lens act together as a compound lens To project an inverted image onto the retina Optic chiasm Optic nerves from both eyes meet and cross at the optic chiasm At the base of the hypothalamus of the brain The information coming from both eyes is combined Then splits according it the visual field Corresponding halves of the field of view (left & right) Sent to the left and right halves of the brain to be processed The right side of primary visual cortex deals with the left half of the field of view from both eyes (similarly for the left brain) Small region in the center of the field of view is processed redundantly by both halves of the brain lgn - - Lateral geniculate nucleus Sensory relay nucleus in the thalamus of the brain Consists of six layers in humans and other primates starting from catarhinias (including apes) One type of ganglion cell (m cell) sends output to the bottom two layers Another type of ganglion cell (p cell) projects to the top four layers Visual information reaching the cortex has been processed by at least four distinct neurons Photoreceptors Bipolar cells Ganglion cells Lgn cells Lgn cells have receptive fields responding If the stimulus falls within a very limited region of space (one degree of visual angle) Layers 1, 4, 6 – correspond to information from the contralateral (crossed) fibers of the nasal retina (temporal visual field) Layers 2, 3, 5 – correspond to information from the ipsilateral (uncrossed) fibers of the temporal retina (nasal visual field) Visual cortex Largest system in the human brain - Responsible for processing the visual image Lies at the rear of the brain above the cerebellum (highlighted in the photo) Primary visual cortex (v1) - region that receives information directly from the lgn V1 - Initial cortical processing area for vision Located in the most posterior portion of the occipital lobe (brodmann area 17) Most studied visual area in the brain In mammals it is located in the posterior pole of the occipital lobe Simplest, earliest cortical visual area Highly specialized for processing information about static and moving objects Excellent in pattern recognition Equivalent to the striate cortex (brodmann area 17 – defined by its anatomical location) Cells in v1 have slightly larger receptive fields This magnification process continues through the visual system Cells in the temporal lobe have receptive fields that may encompass an entire hemifield simple cells mainly in layers 4 and 6 Have distinct excitatory and inhibitory regions Cell that responds primarily to oriented edges and gratings Such cells are tuned to different - frequencies and orientations Calculate edges Complex cells Can be found in the v1, v2, v3 Will respond primarily to oriented edges and gratings However, it has a degree of spatial invariance Means – receptive field cannot be mapped into fixed excitatory and inhibitory zones Some respond to patterns of light in a certain orientation within a large receptive field Regardless of the exact location Some respond optimally only to movement in a certain direction In v2 Use the information from many simple cells to represent corners and edge terminations End-stopped cells Are thought to detect singularities like line and edge crossings, vertices and line endings Neuron in any visual area of the cerebral cortex Maximally responsive to a line of a certain length or to a corner of a larger stimulus Reduced or absent response when the line or corner is extended beyond a certain point Orientation columns Organized regions of neurons Excited by visual line stimuli of varying angles Columns are located in the primary visual cortex (v1) and span multiple cortical layers Neurons with similar properties are arranged in columns perpendicular to the surface of the cortex Range the six cortical layers until they reach the white matter Contralateral projection Between hemisphere and the correspondent side of the body Example: moving the right hand will make the left hemispheres activity go up Upside-down projection 'Images' in your brain are just collections of neural activations, and not actual pictures They cannot have an orientation brain is capable of flipping your visual field if required as measured through perceptual adaptation experiments using inversion glasses Receptive fields Center-surround receptive field Allows ganglion cells to transmit information Whether photoreceptor cells are exposed to light About the differences in firing rates of cells in the center and surround This allows the ganglion cells to transmit information about contrast Size of the receptive field governs the spatial frequency of the information Small receptive fields Stimulated by high spatial frequencies (fine detail) Large receptive fields Stimulated by low spatial frequencies (coarse detail) Retinal ganglion cell receptive fields convey information about: Discontinuities in the distribution of light falling on the retina Often specify the edges of objects - - Retinotopy / retinal mapping / retinally mapped Mapping of visual input from the retina to neurons Particularly those neurons within the visual stream Retinotopy mapping in humans is done with functional magnetic resonance imaging (fmri) Subject inside the fmri machine focuses on a point Then retina – stimulated with a circular image or angled lines about focus point Radial map displays the distance from the center of vision Angular map shows angular location using rays angled about the center of vision Combining both maps – you can see separate regions of the visual cortex Visual pathways Eye to V1, V2 Magnocellular Also called M-cells Neurons located within the Adina magnocellular layer of the lateral geniculate nucleus (LGN) of the thalamus Part of the visual system Characterized by their relatively large size compared to parvocellular cells Receive input from parasol ganglion cells Cannot provide finely detailed or colored information, but still provides useful static, depth and motion information High light/dark contrast detection More sensitive at low spatial frequencies than high spatial frequencies Important for providing information about the location of objects Can detect the orientation and position of objects in space, information that is sent Information = important – detecting the difference in positions of objects on the retina of each eye (important tool in binocular depth perception) Cells in the M pathway – ability to detect high temporal frequencies and can detects quick changes in the positions of an object Parvocellular Also called P-cells Neurons located within the parvocellular layers of the lateral geniculate nucleus (LGN) of the thalamus More modern than M-cells Receive their input from midget cells (type of ganglion cell) (axons of midget cell are exiting the optic tract) Information from each eye is kept separate at this point and continues to be segregated – until processing in the visual cortex Sensitive to color and capable of discriminating fine details than the M cell Greater spatial resolution than M cells Lower temporal resolution than M cells Koniocellular Neuron with a small cell body Located in the koniocellular layer of the LGN (in primates and humans) Quantity of neurons = number of M cells Present between the layers (picture) Koniocellular layers are much thinner due to their size Neurochemically and anatomically distinct from M and. P cells Three proteins Calbindin Alpha subunit of type II calmodulin-dependent protein kinase Gamma subunit of protein kinase C Some cells respond to color Some reacts to achromatic gratings Contribute to brightness contrast information and color contrast in species with color vision - Contribute to eye movement-related signals After V2 Dorsal (‘where’ or ‘how’) Involved in guidance of actions (eg. Reaching) and recognizing where objects are in space Also known as parietal stream, where stream, how stream Stretches from the primary visual cortex (V1) in the occipital lobe forward into the parietal lobe Interconnected with the parallel ventral stream (what stream) Commences with purely visual functions in the occipital lobe Gradually transferring to spatial awareness Termination in parietal lobe Posterior parietal cortex = essential for “the perception and interpretation of spatial relationships, accurate body image, learning of tasks involving coordination of the body in space” Contains individually functioning lobules Lateral intraparietal sulcus (LIP) contains neurons that produce enhanced activation when attention is moved onto the stimulus Ventral intraparietal sulcus (VIP) where visual and somatosensory information are integrated - - Perception Ventral (‘what’) - Associated with object recognition and form representation - Described as “what” stream - Strong connections to The medial temporal lobe (stores long-term memories) The limbic system (controls emotions) Dorsal stream (deals with object locations and motion) - Gets main input from the parvocellular layer of the LGN of the thalamus These neurons project to V1 sublayers 4Cß, 4A, 3B, 2/3a From there the ventral pathway goes through V2 and V4 to areas of the inferior temporal lobe Areas of the inferior temporal lobe - Posterior inferotemporal (PIT) - Central inferotemporal (CIT) - Anterior inferotemporal (AIT) - Each visual area contains a full representation of visual space It contains neurons whose receptive fields together represent the entire visual field Visual information enters the ventral stream through the primary visual cortex Travels through the rest of the areas in sequence All areas in the ventral stream – influenced by extraretinal factors Attention Working memory Salience Damage to ventral stream – inability to recognize faces or interpret facial expressions Perception is a unified whole (binding problem) - Ability of the brain to construct uniform perceptions from a multitude of sensory impressions - In the visual system, structures are known that are activated more strongly by certain shapes, colors or movements in their receptive field than when other patterns are shown - The property of some neurons is the basis of the idea that the sensory information is broken down into such "basic components" - and then put back together again – question? - The binding problem raises the question of how these signals are linked to the overall impression Pathways for visual perception Ventral stream Dorsal stream Computational problems in perception Object constancy Also called perceptual constancy Tendency of animals and humans to see familiar objects as having standard shape, size, color, or location Regardless of changes in the angle of perspective, distance or lighting Impression rends to conform to the object as it is or is assumed to be, rather than to the actual stimulus Responsible for the ability to identify objects under various conditions Seem to be “taken into account” during a process of mental reconstitution of the known image - Reduced by limited experience with the object and by decreasing the number of environmental cues that aid in identification of the object View-invariant recognition Does not happen by simple analysis of the stimulus information The perceptual system extracts structural information about the components of an object and the relationship between these components key to successful recognition is that critical properties remain independent of viewpoint Bicycle example The properties might be features such as an elongated shape running along the long axis, combined with a shorter, stick-like shape coming off of one end With two circular- shaped parts, we could recognize the object as a bicycle from just about any position View-dependent recognition Posit that people have a cornucopia of specific representations in memory Key idea is that the stored representation for recognizing a bicycle from the side is different from the one for recognizing a bicycle viewed from above Our ability to recognize that two stimuli are depicting the same object is assumed to arise at a later stage of processing They seem to place a heavy burden on perceptual memory Each object requires multiple representations in memory, each associated with a different vantage point Shape encoding recognition involve hierarchical representation in which each successive stage adds complexity Simple features such as lines can be combined into … Edges Corners Intersections … which are grouped into parts and the parts grouped into objects (as processing continues up the hierarchy) People recognize a pentagon because it contains five-line segments of equal length (joined together) to form five corners that define an enclosed region (figure 6.12) The same five-line segments can define other objects such as a pyramid Here there are only four points of intersection (not five) and the lines define a more complicated shape that implies – three dimensional Investigate how to encode Identify areas of the brain that are active when comparing contours that form a recognizable shape versus contours that are just jungled grand-mother cell coding Assumption that the final percept of an object is coded by a single cell Cells are constantly firing and refractory Coding scheme of this nature would be highly susceptible to error If a gnostic unit were to die, we would expect to experience a sudden loss for an object Cannot adequately account for how it is possible to perceive novel objects gnostic theory does not account for how the grandmother cell would have to adapt as grandmother changed over time The type of neuron that can recognize a complex object has been called a gnostic unit (from the Greek gnostikos, mean- ing “of knowledge”), referring to the idea that the cell (or cells) signals the presence of a known stimulus—an object, a place, or an animal that has been encountered in the past. Ensemble coding Recognition is not due to one unit but to the collective activation of many units Readily account for why we can recognize similarities between objects and may confuse one visually similar object with another Both objects activate many of the same neurons Losing some units might degrade our ability to recognize an object, but the remaining units might suffice Account for our ability to recognize novel objects Novel objects bear a similarity to familiar things Our percept results from activating units that represent their features Mind reading Encoding - Transform sensory data into a form of mental representation In storage you keep encoded information in memory In retrieval you pull out or use information stored in memory Refers to how you transform a physical, sensory input into a representation that van be placed into memory Encode our memories to store them Short term storage - Short term memory Acoustic code is more important than a visual code Semantics did not matter much for processing Appears to be primarily acoustic May be some secondary visual information than acoustic information Long term storage Most information stored in long term memory primarily is encoded semantically Acoustic information, semantic information, visual information – encoded in long term memory Decoding Decoding techniques interrogate more of the information in the brain scan Rather than asking which brain regions respond strongly to faces They use both strong and war responses to identify subtler pattern of activity These recordings are fed into a “pattern classifier” (computer algorithm that learns the patterns associated with each picture or concept) Once a program has seen enough samples it can start to conclude what the person is looking at or thinking about Pattern recognition Cognitive process that matches information from a stimulus with information retrieved from memory Occurs when information from the environment is received and entered into short-term memory Causing automatic activation of a specific content of long-term memory Super important for humans and animals Koala uses pattern recognition to find and consume eucalyptus leaves Development of neural networks in the outer layer of the brain in humans has allowed for better processing of visual and auditory patterns Six main theories of pattern recognition Template matching Mist basic approach to human pattern recognition Theory – assumes: every perceived object is stored as a “template” into long-term memory Incoming information is compared to these templates to find an exact match Example A a a are all recognized as a but not b This theory – can’t explain how new experiences can be understood without being compared to an internal memory template Prototype-matching Compares incoming sensory input to one average prototype Proposes – exposure to a series of related stimuli leads to the creation of a “typical” prototype based on their shared feature Reduces number of stored templates by standardizing them into a single representation Supports perceptual flexibility – allows for variability in the recognition of novel stimuli Example Suppose a child has never seen a lawn chair before, but given the characteristics of a regular chair, the child would still be able to recognize it as a chair Feature analysis Try to explain how humans are able to recognize patterns in their environment Proposes – nervous system sorts and filters incoming stimuli to allow the human or animal to make sense of the information Proposes an increasing complexity in the relationship between detectors and the perceptual feature When features repeat or occur in a meaningful sequence We are able to identify these patterns because of our feature detection system Recognition-by-components Proposes – humans recognize objects by breaking them down into their basic 3d geometric shapes called geons Example Break down something like a coffee cup We have the hollow cylinder that holds the liquid and a curved handle off the side that allows to hold it Bottom-up Data driven processing Originates with the stimulation of the sensory receptors Claims – perception can be explained solely in term of the environment Stated – sensation is perception and there is no need for extra interpretation as there is enough information in our environment Example Holding a flower at the center of a person’s fields. The sight of the flower and all the information about the stimulus are carried from the retina to the visual cortex in the brain Top-down Refers to the use of background information in pattern recognition Always begins with a person’s previous knowledge and makes predictions due to this already acquired knowledge We construct our perception of reality and these perceptions are hypotheses or propositions based on past experiences and stored information Formation of incorrect propositions will lead to errors of perception – visual illusions Example Given a paragraph written in hard-to-read handwriting Easier to understand if you read the whole paragraph (instead of reading the words in separate terms) Brain may be able to perceive and understand the main idea of the paragraph Due to the context supplied by the surrounding words - - - - Vicarious ai & deep learning deep learning Also known as deep structured learning Applied to fields including: Computer vision Machine vision Speech recognition Natural language processing Audio recognition Social network filtering Etc. “deep” because of the use of multiple layers in the network Part of a broader family of machine learning methods Based on artificial neural networks with representation learning Learning can be supervised, semi-supervised or unsupervised Supervised Machine learning a function that maps an input to an output based on example input and output pairs Semi-supervised Approach to machine learning that combines a small amount of unlabeled data (can produce considerable improvement in learning accuracy) during training Unsupervised Type of machine learning that looks for previously undetected patterns in a data set with no pre-existing labels (minimum human supervision) memory - Storage of retrievable information, outcome of learning Encoding Processing if new information Two types of encoding Acquisition Sustaining some of the sensory stimuli to enter into memory Sensory systems are constantly being bombarded by tons of stimuli Most only produce a very brief transient sensory response hat fade quickly (1000 ms after representation) During this period the stimuli are available for processing State called a sensory buffer Only some of these stimuli are sustained and make the cut into short term memory (acquisition) Consolidation Stabilize a memory over time resulting in long term memory This can occur over days to months even years Creates a stringer representation over time Storage Represents the permanent record of the information Result of acquisition and consolidation Retrieval Accessing stored information Using it to create a conscious representation or to execute a learned behavior such as a motor act anatomy of memory Hippocampus Embedded deep into the temporal lobe Major role in learning and memory Part of the limbic system (which regulated emotions) Processes long-term memory and emotional responses If this is being removed Severe anterograde amnesia of declarative memory Learn new skills but not remember learning Essential for memory consolidation Active when new information is encoded Also involved in the retrieval if information from long-term memory Place cells Pyramidal neuron within the hippocampus Becomes active when entering a particular place in its environment Act as a cognitive representation of a specific location in space known as a cognitive map Provide evidence that the hippocampus has cells that encode contextual information Entorhinal cortex Area of the brain located in the medial temporal lobe Functions as a hub in a widespread network for memory, navigation and the perception of time Main interface between the hippocampus and neocortex Plays an important role in declarative (autobiographical/episodic/semantic) memories and in particular spatial memories Memory formation, memory consolidation, and memory optimization in sleep Also responsible for the pre-processing (familiarity) of the input signals Association of impulses from the eye and the ear occurs in the entorhinal cortex Grid cells Type of neuron within the entorhinal cortex Fires at regular intervals as an animal navigates an open area allowing it to understand its position in space by storing and integrating information about Location Distance Direction Have been found in animals (rats, mice, bats, monkeys) and humans Unique properties - have firing fields over the entire environment (place fields are restricted to certain specific regions of the environment) firing fields are organized into hexagonal Hexagonal lattice lattice Firing fields are generally equally spaced apart such that the distance from one firing field to all six adjacent firing fields is approximately the same Firing fields are equally positioned Such that six neighboring fields are located at approximately 60 degree increments Standard consolidation theory (squire) memory consolidated fast in the hippocampus Via entorhinal cortex During sleep consolidated in other parts of cortex Proposed by larry squire and his colleagues Considers the neocortex to be crucial for the storage of fully consolidated long-term memories Considers - the hippocampus plays a temporary role In this view: The representation of an event that are distributed throughout the cortex come together in the medial temporal lobe The hippocampus binds them Then through some sort of interaction between the medial temporal lobe and the neocortex, the bound information – slowly transferred and replaced by a permanent memory trace (in the neocortex) Occurs after repeated reactivation of the memory, creates direct connections within the cortex between the various representations Process takes place when being asleep or awake Eventually makes the memory independent of the hippocampus Key Points Memory and Anatomy of Memory Learning is the process of acquiring new information, the outcome of which is memory Learning and memory have many stages including encoding, storage and retrieval What is known as the medial temporal lobe memory system is made up of the hippocampus and the surrounding rhinal and parahippocampal cortices Other areas involved with memory include the prefrontal cortex, the parietal cortex and subcortical structures The hippocampus is critical for the formation of long- term memory. Cortex surrounding the hippocampus is critical for normal hippocampal function in memory. The delayed non-match to sample task is used to assess memory in non-human primates. The amygdala is not a crucial part of the system for epi- sodic memory, but it is important for emotional memory. Neurons that activate when rats are in a particular place and facing a particular direction have been identified in the hippocampus and are called place cells. They provide evidence that the hippocampus has cells that encode contextual information. types of memory Long term memory Declarative (explicit) memory Defined as memory for events and for facts, both personal and general, that we have conscious access to and that can be verbally reported Dependent on the medial temporal lobe Can be broken down into two memories episodic memory Memories of personal experience That we recall about our own lives and what, where, when and with whom they happened They differ from personal knowledge Example (1) You have personal knowledge of what day you were born but you do not remember the experience Always include the self as the agent or recipient of some action Example (2) The memory of falling off your new red bi- cycle (what) on christmas day (when), badly skinning your elbow on the asphalt driveway (where), and your mother (who) running over to comfort you is an episodic memory The result of rapid associative learning in that the what, where, when, and who of a single episode - Semantic memory Is objective knowledge that is factual in nature but does not include the con- text in which it was learned Example (1) You may know that corn is grown in iowa, but you most likely don’t re- member when or where you learned that fact Fact can be learned after a single episode, but it may take many exposures Reflects knowing facts and concepts such as how to tell time, who the lead guitarist is for the rolling stones, and what quantum mechanics is all about Nondeclarative (implicit) memory It cannot be “declared,” that is, verbally reported Also known as implicit memory Knowledge that we have no conscious access to Several types of memory fall under this category: priming, simple learned behaviors that derive from conditioning, habituation, sensitization, and procedural memory, such as learning a motor or cognitive skill This form of memory is revealed when previous experiences facilitate performance on a task that does not require intentional recollection of the experiences Is not dependent on the medial temporal lobe Involves other brain structures, including the basal ganglia, the cerebellum, the amygdala, and the neocortex Procedural memory One form of nondeclarative memory Depends on extensive and repeated experience Tasks that require us to use procedural memory include learning motor skills like how to ride a bike, type, or swim, and learning cognitive skills such as how to read One test of procedural learning is the serial reaction time task participants sit at a console having four buttons Placing the fingers of one hand over the buttons, participants would press buttons that correspond to locations of stimuli in front of them Each button corresponds to one of four lights The mapping between button and light can simply be their spatial relationships The task would be to press the button with the finger that corresponds to the light that is illuminated The lights can be flashed in different sequence Over time, normal participants respond faster to the repeating sequence than they do to a totally random sequence Their improved performance indicates that they have learned the sequence When asked whether the sequences were random, however, participants report that the sequences were completely random Do not seem to know that any pattern existed, yet they learned the skill Such behavior is typical of procedural learning, which requires no explicit knowledge about what was learned Patients with disorders of the basal ganglia or inputs to these subcortical structures show poor performance on a variety of procedural learning tasks Priming Refers to a change in the response to a stimulus, or in the ability to identify a stimulus, following prior exposure to that stimulus Example (1) If you were to see a picture of bicycle handlebars from an odd angle, you would recognize them as part of a bike faster if you had just seen a typical picture of a bike Priming can be perceptual, conceptual, or semantic. Acts within the perceptual representation system (PRS) The structure and form of objects and words can be primed by prior experience, and the effects persist for months Involves stimuli that have similar forms Example The word "goat" will evoke a faster response when it is preceded by the word "boat" because the two words are perceptually similar. Conceptual priming Involves a stimulus and response that are conceptually related Example Words such as "desk" and "chair" are likely to show priming effects because they are in the same conceptual category. - Semantic priming Involves words that are associated in a logical or linguistic way Example Responding to the word "banana" more rapidly after being primed with the word "yellow" is an example of semantic priming. Short term memory Limited capacity Hold information for longer than sensory stores (30s) Fragile: distraction causes you to forget Two types Sensory memory Information decays very rapidly (within 0.5 seconds) Information can be erased Stored in sensory memory just long enough to be transferred to short-term memory Humans have five traditional senses: sight, hearing, taste, smell, touch Sensory memory allows individuals to retain impressions of sensory information after the original stimulus has ceased The formation of a sensory memory trace is only weakly dependent on attention to the stimulus Iconic memory For visual was the first sensory store to be investigated with experiments dating back as far as 1740 iconic memory in humans has a large capacity, but decays very rapidly Echoic memory For auditory Auditory information travels as sound waves which are sensed by hair cells in the ears Information is sent to and processed in the temporal lobe holds information for 2–3 seconds to allow for proper processing regards to language, a characteristic of children who begin speaking late in development is reduced duration of echoic memory Echoic memory is a fast-decaying store of auditory information In the case of damage to or lesions developing on the frontal lobe, parietal lobe, or hippocampus, echoic memory will likely be shortened and/or have a slower reaction time Working memory Cognitive system with a limited capacity that can hold information temporarily important for reasoning and the guidance of decision-making and behavior Often used synonymously with short-term memory, but some theorists consider the two forms of memory distinct Assuming that working memory allows for the manipulation of stored information Widely acknowledged as having limited capacity Represents a limited-capacity store for retaining information over the short term (maintenance) and for performing mental operations on the contents of this store (manipulation) Example We can remember a list of numbers, and we can also add (manipulate) them in our head by using working memory Atkinson & shiffrin’s modal (or multi-store) model Proposes that information is first stored in sensory memory From there, items selected by attentional processes can move into short-term storage Once in short-term memory, if the item is rehearsed, it can be moved into long-term memory Suggests that, at each stage, information can be lost by Decay (information degrades and is lost over time) Interference (new information displaces old information) Or a combination of the two Formalized the idea that discrete stages of memory exist and that they have different characteristics Strong serial structure Information coming into sensory memory can be passed to short-term memory and only then into longterm memory Baddeley’s working memory model Argued that the idea of a unitary short-term memory was insufficient to explain the maintenance and processing of information over short periods Proposed a three-part working memory system consisting of a central executive mechanism for controlling two subordinate systems - - - Phonological loop Hypothesized mechanism for acoustically coding information in working memory Studies that asked participants to recall strings of consonants Letters were presented visually, but the pattern of recall errors indicated that perhaps the letters were not coded visually over the short term Participants were apparently using an acoustic code More likely to replace a presented letter with an erroneous letter having a similar sound (e.g., t for g) Rather than one with a similar shape (e.g., q for g) Might have two parts Short-lived acoustic store for sound inputs Articulatory component that plays a part in the subvocal rehearsal of visually presented items to be remembered over the short term Visuospatial sketchpad Short-term memory store that parallels the phonological loop and permits information storage in either purely visual or visuospatial codes Studies of participants who were instructed to remember a list of words using either A verbal strategy such as rote rehearsal Or A visuospatial strategy based on an imagery mnemonic Participants were better on the memory test when they used the visuospatial strategy Verbal strategy, however, proved better when the participants were required to concurrently track a moving stimulus by operating a stylus during the retention interval Free recall experiments Show serial position curve Primacy effect Superior recall of item near begin Is the beginning You remember it because that is where you started For example A subject who reads a sufficiently long list of words is more likely to remember words toward the beginning than words in the middle Recency effect Superior recall of item near end Is the finish You remember the end the best Is increased when too much information is presented too quickly and is reduced when coupled with other tasks With respect to jury memory, allowing note taking could also reduce it Constructive memory Prior experience Has an influence on memory This is a problem with eye witnesses How fast were the cars going when they … each other? Smashed, collided, bumped, hit, contacted Encoding specificity State, mood and context have influence on memory Memory performance is optimal when there is a match in processes/context between learning and retrieving Flashbulb memories Where were you on 9/11 Where were you when you heard trump was elected president of the USA Emotion Memory deficits Amnesia Anterograde For events after the injury Loss of ability to form new memories No consolidation Loss of memory for events that occur after a lesion Inability to learn new things Retrograde For events before the injury Loss of memory for events that happened in the past No retrieval Sometimes retrograde amnesia is temporally limited, extending back only a few minutes or hours Ribot’s law More loss of the most recent events (closest to the injury) in retrograde amnesia First postulated by Théodule Ribot (French psychologist) Amnesia can differentially affect short term memory, working memory or long-term memory abilities - Alzheimer’s disease Amyloid plaques and neurofibrillary tangles in brain Memory impairment and later general impairment Starts with episodic memory Starts with entorhinal cortex - Largest risk factor: age (6%>80 y ) Hebbian learning Associative learning Cells that fire together wire together Cells that fire together strengthen synapses Cells that fire separately loose synapse Asserts that if a neuron is simultaneously Little Dictionary: activated by a pathway with a weak input and another pathway with a strong input LTP – long term potentiation both pathways show LTP and the weak (potentiate = to strengthen) synapse becomes stronger When two weak inputs (W1 and W2) and one LTD – long term depression strong input (S1) are given to the same cell When W1 and S1 are active EPSP – excitatory postsynaptic together, W1 is strengthened but potentials W2 is not If W2 and S1 are active together, W1 is not affected by the LTP induced from W2 and S1 Hebb’s rule / Hebb’s law If a synapse is active when a postsynaptic neuron is active, the synapse will be strengthened LTP - - (E)PSP is increased after presynaptic stimulation Long term synapse strengthening Occur in the other two excitatory projection pathways of the hippocampus Last days or weeks in living animals Varies in the three pathways Also takes place in other places of the brain Amygdala Basal ganglia Cerebellum Cortex Can be recorded by placing stimulating electrodes on the performant pathway and a recording electrode on a granule cell of the dentate gyrus - A single pulse is presented and the resulting EPSP is measured - Size of the first recording is the strength of the connection before the LTP is induced - Then the performant pathway is stimulated with a burst of pulses - After LTP is induced, a single pulse is sent again and the magnitude of the EPSP in the postsynaptic cell is measured - Magnitude of EPSP increases after LTP is induced - Associative LTP – extension of the Hebbs Law LTD - Long term depression When the pulses are presented slowly – low frequency pulses – LTD develops (E)PSP is decreased after presynaptic stimulation Long term synapse weakening Occurs at synapses in cerebellar purkinje neurons which receive two forms of excitatory input One from a single climbing fiber and one from hundreds of thousands of parallel fibers Decreases the efficacy of parallel fiber synapse transmission - LTD has been observed in the visual cortex Long Term Depression Proposed to be involved in ocular dominance Reduces PSP Types of Learning Supervised learning Teacher Important issues Where do you get examples with answers? Which examples are important? How do you translate an error into a correct answer next time so that it generalizes? Nature (school, university) Machines (Articial neural networks (ANN), support vector machines (SVM)) Reinforcement learning A bit of help (rewards) Important issues What is the value of actions for future rewards? Nature (cookie when you are good) Machines (Q learning) Unsupervised learning All on your own Important issues What is shared structure and what is unique? Principle component analysis (PCA) Content addressable memories (CAM) Learning over different timescales Developmental learning Perception level just before and after birth Humans: weeks to months Shape basic connections in the brain/machine Learn to process basic stimuli from the environment Learn invariants A lot of (but not exclusively) unsupervised learning: Artificial Neural Network Behavioral learning At the task level throughout life Humans: years Shape more complex/advanced connections in the brain/machine Learn which cause leads to reward at a later time Learn credit assignment A lot of (but not exclusively) reinforcement learning Evolutionary learning At the architecture level over generations Humans: centuries/millennia Shape the basic architecture of the brain/machine Learn which brain/machine architecture is fittest I.e. Does the best developmental learning and behavioral learning Natural selection, genetic algorithms Artificial Neural Networks Computing systems vaguely inspired by the biological neural networks that constitute animal brains A very crude approximation of biological neural networks They consist of nodes (neurons, units, …) A node has input/output They consist of connection between node Connections have a weight (synaptic strength) Learning is changing the connection weights The input function is (usually) defined as: in = ∑ W *a = W ·a i j j,i k i i This linear weighted sum is transformed into an activation value by an non-linear activation function: Step function Sign function Sigmoid function - Each connection, like the synapses in a biological brain, can transmit a signal to other neurons artificial neuron that receives a signal then processes it and can signal neurons connected to it Node/Unit/Neuron Receives input from some other nodes, or from an external source and computes an output Each input has an associated weight (w), which is assigned on the basis of its relative importance to other inputs Applies a function f (defined below) to the weighted sum of its inputs Types of general structure Fully connected All units to all Are a type of artificial neural network where the architecture is such that all the nodes or neurons In one layer are connected to the neurons in the next layer Feed-forward One layer to next Wherein connections between the nodes do not form a cycle Was the first and simplest type of artificial neural network The information moves in only one direction (forward) From the input nodes Through the hidden nodes (if any) And to the output nodes No cycles or loops in the network Recurrent Some back connections Connections between nodes form a directed graph along a temporal sequence Allows it to exhibit temporal dynamic behavior Perceptron - Has an input layer and an output layer Uses the step or sign function Single-layer, feed-forward network Because of the step or sign function: binary activation Supervised learning rule for weight changes Learning in the Perceptron: Algorithm in words Calculate the error What values can this error take, assuming a step function) If error = 0 then stop Else adjust the weights Adjusts weights only if input !=0 Increase weight if error=1 Decrease if error = -1 Perceptron Algorithm: If error (target output – current output) != 0 then Weight change = (some small constant) * (error) * (input) In one go: Weight change = (some small constant) * (target output – current output= * (input) Perceptron weight learning rule ∆w = μ(t -a )a = μδ a ki k k i k i ∆w = weight change to node k from node i ki a = is output of node i, a of node k i k t = target output for node k k δ = error for node k k μ = learning constant (usually between [0,1]) The Hebbian weight learning rule: - ∆w = μa a ki k i Linear separability - Multi-layer perceptron Multi-layer perceptron were hypothesized in the 50’s There is an obvious problem however We cannot use our Perceptron learning rule to update the nodes in the hidden layer Why? We do not know the target output for the hidden notes - - Back-propagation learning (‘back-prop’) Back-prop can be generalized to any number of layers It can be shown that the perceptron/backprop learning rule is completely identical to an algorithm called gradient descent (aka hill-climbing or steepest-descent) Disadvantages: Learning might take a long time, and is often depended on initial Weights (slow and inefficient) It is a black-box New(online)learning will overwrite old representations Cannot find the globally optimal solution (local minimum) It can be over trained: does not generalize to new examples well Deep learning Many layers Back-prop Convolutional deep neural nets Analogous to human visual system Local connectivity in convolutional layers Often pre-sets, pre-trained or from unsupervised learning Fully connected (dense) higher layers Supervised learning Really good image recognition About as good as humans More and more complex representations in the NN layers Functions of Attention Vigilance Signal detection Try to detect whether we sensed a signal to take (swift) action Search Actively looking for a signal Selective attention Actively focus on some information and ignoring other information Divided attention Perform more than one task simultaneously and shift attention between them Attention is the ability to focus awareness on one stimulus, thought or action while ignoring other irrelevant stimuli, thought or actions Signal detection theory Provides a precise language and graphic notation for analyzing decision making in presence of uncertainty Offers a way to analyze many different kinds of decision problems Sensitivity Many hits Few false alarms Specificity Many correct rejection Few misses Hit, Miss, False alarm, Correct rejection Explained in the picture In this case if the tumor is absent and the doctor responds yes = false alarm Search - In everyday perception, voluntary attention (driven by our goals) and reflexive attention (driven by stimuli in the world) Interact in a push-pull fashion, struggling to control the focus of our attention Feature (pop-out) search - - One set of experiments They observed that targets are located more quickly among a field of distracters if the target can be identified by a single stimulus feature, such as color Example A red O among green Xs and Os Doesn’t matter how many distracters appear in the array Can demonstrate this relation by plotting participants’ reaction times as a function of the number of distracter items in the display Conjunction search Proposed that while elementary stimulus features such as color, motion, shape, and spatial frequency Can be analyzed pre-attentively and in parallel within multiple specialized feature maps Spatial attention is more complicated Spatial attention must be directed to relevant stimuli in order to integrate the features into the perceived object Necessary to link the information in the different feature maps so that the target can be analyzed and identified Feature integration theory A psychological theory of visual perception Based on the idea that the visual system can process in parallel elementary features Such as color, shape, and motion Requires spatial attention to bind the features that define an object Spotlight of attention must move sequentially from one item in the array to another Similarity theory Suggests that attention is not drawn to locations but rather to image objects Search efficiency depends on similarities Between objects in the scene Possible targets Between objects within the scene Guided search theory Model of visual attention Explains how we find out intended target when looking in a crowded visual field First stage, basic stimuli features are processed simultaneously at all locations across the visual field Second, limited-capacity serial stage, processing is more complex and restricted to a particular item or location based on information obtained in the previous stage Selective Attention Cocktail party problem/effect Imagine yourself at a Super Bowl party having a conversation with a friend How can you focus on this single conversation while the TV is blasting and boisterous conversations are going on around you? Psychologist E. C. Cherry (1953) wondered the same thing while attending cocktail parties Selective auditory attention allows you to have a conversation - bar or party - while ignoring the rest of the sounds around you Selectively attending You can perceive the signal of interest amid the other noises If the person you are conversing with is boring You can give covert attention to a conversation going on behind you While still seeming to focus on the conversation in front of you Dichotic listening task Investigated this ability by designing a cock- tail party in the lab Normal participants, wearing head- phones, listened to competing speech inputs to the two ears This setup is referred to as dichotic listening Then asked the participants to attend to and verbally “shadow” the speech Coming into one ear, while simultaneously ignoring the input to the other ear Discovered that under such conditions, participants could not Report any details of the speech in the unattended ear In fact, all they could reliably report from the unattended ear was whether the speaker was male or female Attention, in this case voluntary attention, affected what was processed propose that attention to one ear results in better encoding of the inputs to the attended ear - loss or degradation of the unattended inputs to the other ear Posner cuing task Have suggested that this attentional spotlight affected reaction times by influencing sensory and perceptual processing The stimuli that appeared in an attended location were processed faster Than the stimuli that appeared in the unattended location Enhancement of attended stimuli, a type of early selection Suggests that changes in perceptual processing can happen When the participant is attending a stimulus location Voluntary attention is our ability to intentionally attend to some- thing, such as this book a goal-driven process, meaning that goals, knowledge, or expectations used to guide information processing Reflexive attention bottom-up, stimulus-driven process in which a sensory event maybe a loud bang, the sting of a mosquito, a whiff of garlic, a flash of light or motion captures our attention these two forms of attention differ in their properties and perhaps partly in their neural mechanisms Overt /covert attention when you turn your head to orient toward a stimulus whether it is for your eyes to get a better look your ears to pick up a whisper your nose to sniff the frying bacon you are exhibiting overt attention Key Points Attention involves both top-down (voluntary), goal-directed processes and bottom-up (reflexive), stimulus-driven mechanisms. Attention can be either overt or covert. According to early-selection models, a stimulus need not be completely perceptually analyzed before it can be selected for further processing or rejected as irrelevant. Broadbent proposed such a model of attention. Late-selection models hypothesize that attended and ignored inputs are processed equivalently by the perceptual system, reaching a stage of semantic (meaning) encoding and analysis where selection may occur. Our perceptual system contains stages at which it can process only a certain amount of information at any given time, what are called limited-capacity stages which result in processing bottlenecks. Attention limits the information to only the most relevant, thereby preventing overload of the limited-capacity stages. Cuing tasks, where the focus of attention is manipulated by the information in the cue, are often used to study the effect of attention on information processing. Spatial attention is often thought of metaphorically as a “spotlight” of attention that can move around as the person consciously desires, or that can be reflexively attracted by salient sensory events. Anatomy of attention attention system uses subcortical and cortical networks within the brain Interact to enable us to selectively process information in the brain Several subcortical structures are relevant to both attentional control and selection Superior colliculus in the midbrain and the pulvinar Involved in aspects of the control of attention In the cortex – several areas that are important to attention Frontal cortex Posterior parietal cortex Posterior superior temporal cortex Also include some medial brain structures Anterior cingulate cortex Posterior cingulate cortex Insula Dorsal Frontoparietal attention network Goal directed driven control For unattended salient stimuli Strongly lateralized to the right Inferior (ventral) parietal and frontal cortex Ventral Frontoparietal attention network Stimulus lateralized to the right Inferior (ventral) parietal and frontal cortex (NOT THE SAME AS DORSAL AND VENTRAL PATHWAYS) Attention deficits ADHD Attention deficit hyperactivity disorder Characterized by Attention deficits (distractibility) Hyperactivity (fidgetiness) Impulsiveness Mood swings Short temper High sensitivity to stress Impaired ability to make and follow plans Have occupational difficulties or antisocial behaviors in adulthood Careful measurements of ADHD people’s behavior help clarify our descriptions of the condition Measurements also enable researchers to quantify improvements that might occur after treatment People with ADHD have difficulty inhibiting their behaviors ADHD often runs in families, and twin studies suggest fairly high heritability Attention to a stimulus is almost synonymous with being conscious of it Attention or consciousness relates to increased brain activity in the areas responsive to a stimulus Damage to parts of the right hemisphere produce spatial neglect for the left side of the body or the left side of objects Sensory neglect results from a deficit in attention, not sensation Common diagnosis in the United States Probably the result of a number of genes as well as environmental influences Many people with ADHD have mild brain abnormalities, but the pattern is small and inconsistent Change blindness Process of failing to notice a change in some scene, usually because the change occurred gradually or while the viewer was moving or blinking his or her eyes If part of a complex scene changes slowly, you probably won’t notice it You might also fail to notice a quick change that occurs while you are blinking your eyes or moving them Quickest way to get people to notice the change is to instruct them where to direct their attention within the scene Change blindness is the process of failing to notice a change in some scene, usually because the change occurred gradually or while the viewer was moving or blinking his or her eyes Neglect - Opposite of attention is inattention, or neglect Right now, presumably you are attending to what you are reading, ignoring what you see above, below, or to the left or right of this page People with damage to parts of the right hemisphere Tendency to ignore the left side of the body and its surroundings or the left side of objects Damage in the left hemisphere does not pro- duce much neglect of the right side Also, generally ignore much of what they hear in the left ear and feel in the left hand, especially if they simultaneously feel something in the right hand Exact location of the damage within the right hemisphere varies, as do the details of what the person neglects Damage to the inferior part of the right parietal cortex tend to neglect everything to the left of their own body Damage to the superior temporal cortex neglect the left side of objects, regardless of their location When a patient with neglect sees a large letter com- posed of small letters, he or she can identify the large letter, even though neglecting part of it when asked to cross off all the small letters. Also, some- one who neglects the left hand pays attention to it when it is crossed over the right hand Automatic and controlled processes Automatization Controlled processes can become automatic processes Example Learning to drive a car Can take a long time Practice effects Rate of learning slows The first few practice sessions – much is learned Over time each session results in less learning Stroop effect Delay in reaction time between congruent and incongruent stimuli Used to create a psychological test Widely used in clinical practice and investigation - Basic task that demonstrates this effect Occurs when there is a mismatch between the name of a color "blue", "green", or "red" And the color it is printed on The word "red" printed in blue ink instead of red ink When asked to name the color of the word it takes longer and is more prone to errors when the color of the ink does not match the name of the color Attentive robots Example BeoBot Attentional SLAM Autonomous city explorer Robots that use a computational attention system to direct their focus of attention to parts of the sensory input which are currently of most potential interest Social robots Autonomous robot Interacts and communicates with humans or other physical agents By following social behaviors and rules attached to its role Research design Correlational - Just measure variables as they occur Used to determine if there is an association You can never infer a causal relationship - Involves a manipulation of one variable Keep other variables constant Can show a causal relationship Experimental Computer modeling - Simulate models of cognition inside the computer Use them to generate testable prediction Most well-known models are based on connectionist approach (ANNs) As computers become more powerful, so do the models Great as a testbed for studying theories about human cognition But: Models always simplify George P. Box: “All Models are wrong, but some are useful” Some models are not biological plausible Some models are too complex they fit almost any data, and can generate almost any desired behavior, but generalize poorly Always many different models that can generate can same behavior John von Neumann: “Give me 4 degrees of freedom and I will fit you and elephant, give me 5 and I will have it wave its trunk at you” Symbolic modeling Often serial and abstract processing Example Expert Systems Connectionism Parallel and distributed processing Example ANN’s Still both at Marr’s algorithmic level Realistic neuronal modeling IBM Blue Brain The EU Human brain project Much closer to Marr’s implementational level Eventually: neuromorphic computing Braitenberg vehicles We see that there is no difference in how the two models process information, but only a difference in their patterns of connectivity The point of Braitenberg’s example is not to model a behavior; rather, it represents how a single computational change – from crossed to uncrossed wiring – can yield a major behavior change. When interpreting such a behavioral difference, we might postulate extensive internal operations and representations Pro’s and Con’s Advantages Can be safer and cheaper than the real world. Able to test a product or system works before building it - Can use it to find unexpected problems Able to explore ‘what if…’ questions Can speed things up or slow them down to see changes over long or short periods of time Disadvantages Mistakes may be made in the programming or rules of the simulation or model The cost of a simulation model can be high - The cost of running several different simulations may be high - Time may be needed to make sense of the results - People’s reactions to the model or simulation might not be realistic or reliable Lesion Studies - - - - Structural Imaging CT / CAT - Provided key insights into the relationship between brain and behavior Study of damage to the human brain allows us to understand functions of certain parts If something is damaged, the changes in behavior are often linked to the damaged region Fundamental concepts Left hemisphere dominant role in language The dependence of visual functions on posterior cortical regions Careful with interpretation of results Which functional area Can have an effect on other regions It is not a normal, healthy brain… Dissociations No dissociation One lesion impairs A and B Example Lesion in parietal cortex impairs visual attention and speech perception Single dissociation One lesion impairs A and B, other only B Example Lesion in parietal cortex impairs visual attention and speech perception And lesion in temporal cortex impairs only speech perception Double dissociation One lesion impairs only A, other lesion only B Example Lesion in parietal cortex impairs only visual attention And lesion in temporal cortex impairs only speech perception Pro’s and Con’s Pros gives inside to damage Cons Brain is complex Damage to one part could affect another Skewing results Results in a non-healthy brain Not a dependable source of research CT = Computed Tomography Uses X-Rays Different tissues absorb different amounts of X-Rays Shows contrast between tissue Is radioactive Causes damage Not always feasible for prolonged study Structural MRI MRI = Magnetic Resonance Imagining MRI system Causes spinning atoms in the field to align to magnetic North/South Then, a radio wave is beamed into the field, causing unmatched atoms to align in opposite directions When the radio wave is turned off, atoms realign, releasing energy We can measure this energy Magnet (1.5T – 9.4T) Higher number = higher magnetic field Why higher fields? The higher the rating, the higher the magnetic field Which results in more accurate readings Gradient coils Creates a secondary magnetic field In a predictable manner Allows us to alter the resonant frequency of certain atoms Radio frequency coil Beams radio waves into the field Protons / spins Spinning atoms What we measure in an MRI Larmour frequency The frequency of the radio pulse Determined by gyromagnetic ratio: Hz/Tesla Gyromagnetic ratio Hertz/Tesla T1 weighted, T2 weighted, T2* weighted Different relaxation times (of the spins realigning) Give different contrast, enhancing images Diffusion MRI / DTI Uses the diffusion of water molecules To enhance contrast in MR Images Anistropic diffusion Directional diffusion within the brain Useful for determining structures in the brain that restrict the flow of water Such as myelinated axons of nerve cells (affected by multiple sclerosis) Pro’s - Gives us increased clarity in the flow of water throughout the brain - Doesn’t tell us much about injuries/lesions Con’s Functional Imaging Single cell recording Famous experiment by Hubel and Wiesel. Cells in the V1 (see visual cortex) respond in a specific manner Pro’s Gives very high resolution Con’s Highly invasive Does not give the whole picture EEG - EEG = Electroencephalogram Measures overall brain activity over time High temporal resolution Uses electrodes to measure activity of groups of neurons that fire in parallel ERP ERP = Event – related potential Once an event occurs, we can measure the brain activity as it happens ECoG Electrodes are placed directly on the surface of the brain Much more invasive, but less noisy Pro’s Very cheap Very high temporal resolution Con’s Very poor spatial resolution Very noisy Impossible to detect MEG - Magnetoencephalogram Measures same activity as EEG Less mobile More accurate in location Pro’s More accurate in spatial resolution (contrast) Con’s Less mobile, less practical Time-frequency analysis We don’t always see everything Do the same thing over and over again Average results - Positron Emission Topography Measures brain activity through a tracer Substance given to patient Sits in bloodstream Subtraction method Allows us to determine very specific locations in the brain Using subtraction of a control vs affected image Pro’s You can measure what you trace depending on the substance given Con’s Radioactive substance must be imbibed Has been replaced by better methods Experiments are short (radioactive decay) Very expansive - fMRI = functional MRI goes beyond subtraction method relies on the general linear model PET fMRI BOLD, oxyHb and deoxyHb Oxygenated vs Deoxygenated Hemoglobin (Hb) has different resonant magnetic frequencies BOLD = blood Oxygen Level Dependent responses Gives us an insight to mixture of blood flow/volume in brain Block design & event-related design Create an event and measure the response over time (2-3 seconds) Rt-fMRI, BCI & Neurofeedback Real-Time fMRI, record results as they happen BCI = Brain Computer Interface, allows us to control computers via thought Neurofeedback feeds brain activity back to the subject. Subject can “train” their thoughts to control devices via BCI Pro’s - Very high spatial resolution Relatively high temporal resolution Non-invasive - Temporally limited - Con’s Spatial & temporal resolution Spatial = location Temporal = time Relative resolutions of modalities Log size Log time plot Methods to perturb function Perturbing function is altering the function of the brain, whether through drugs, stimulants, or otherwise Because damage to the brain does not necessarily tell you what the function of the damaged portion does Removing the spark plug from an engine does the same thing as running out of fuel, but they’re two different things entirely Focusing on more localized, controlled “perturbation” Pharmacological studies There are certain studies that introduce certain types of chemicals, or drugs, known as agonists Which act like neurotransmitters and mimic/increase neurotransmission Antagonists Which bind to cell receptors and block or somewhat dampen neurotransmission) These studies allow us to study the direct effects of certain neurotransmitters or other chemicals on cognitive function TMS - Transcranial magnetic stimulation A large coil generating a strong magnetic field is placed on the skull and causes certain neurons to fire The mechanism is poorly understood, though the applications are many Pulsed tms Pulsing the magnetic field Able to time any given stimulus with the tms pulse to prevent certain actions from happening, such as pulsing in the visual cortex between 70-130ms after the subject is shown a certain letter in the alphabet, preventing them from identifying and naming the letter Repetitive tms Performing tms to specific regions of the brain repeatedly over many sessions Allows certain patients with physiological or psychological disorders to experience temporary relief The more often and prolonged a session is, the longer the residual benefit tDCS - Transcranial direct current stimulation An anode and cathode are placed onto the scalp Neurons underneath these terminals become depolarized Allowing them to become much more excitable and therefore more likely to trigger an ap when a specific stimulus occurs Genetic manipulation One of the greatest scientific challenges of today is in the manipulation of genes To determine the eventual outcome of human development The human genome project was able to successfully map the entire human genome The challenge lies within decoding all of the information, and determine which genes lead to which type of development Such as which genes lead to the development of brain tissue vs muscle tissue, etc Knockout procedures A type of experiment on animals where scientists “knock out” a specific gene Let the specimen develop otherwise normally to see if they can’t detect what changes their manipulation leads to Example Certain types of mice with “knocked-out genes” were unable to develop their cerebellum fully, leading them to develop poor motor control Language & Linguistics Language is uniquely human There is a lack of an upper bound on the number of grammatical sentences in a language Language is recursive Language structure is the same all over the world Human language acquisition is the same all over the world Language is learned from imperfect environments Grammar Refers to the structural rules that govern the composition of words, phrases and sentences in a particular natural language Syntax - Rules that tell us how to organize words (grammar) Phoneme Smallest unit of speech Grapheme Smallest unit of written language Semantics Meaning of language Semantic relationships between words are ab organizational principle of the mental lexicon Morpheme Smallest unit with meaning Lexicon - Mental store of word information Model for the lexicon: semantic network Words and their associative connections Strongly semantically or associatively related words are close Semantically related: similarly colored Associatively related: connected Prosody - Rhythm and pitch of speech Pragmatics Knowledge of social rules Language acquisition Language is special to humans Animals communicate, but no advanced language We learn language despite (motherese) Motherse = simplified and repetitive type of speech, with exaggerated intonation and rhythm (often used by adults when speaking to babies Human language acquisition is the same all over the world Stages Cooing Babbling (6 months of age) One- word utterances (1 year of age) Two-word utterances (1-3 years of age) Basic adult structure Overextension error Use of wrong semantic word Example Everything with four legs is a doggie Irregular verb errors They first get it right: sing – sang Later on, they get it wrong: sing – singed (overgeneralization) Eventually they get it right again: sing – sang Irregular past tense is acquired associatively and regular as rule based Critical period There seems to be a critical period for language acquisition As you get older, you may learn a language, but never with the same mastery as you did when you were younger Anatomy of Language Lateralization Language is left lateralized Left peri-sylvian language network Peri-sylvian language network Inferior frontal cortex Broca’s area Inferior parietal lobule Superior temporal gyrus Wernicke’s area Arcuate fasciculus Aphasia - Brain damage in the language areas often leads to Aphasia Deficits in language comprehension and production Even though articulatory mechanisms are intact Very common after a stroke (40%) - Almost all strokes of the left hemisphere lead to transient aphasia Broca’s aphasia - Also called anterior aphasia or expressive aphasia - Damage of the Broca’s area - Mainly deficits in speech production - Also, comprehension deficits related to syntax (agrammatic aphasia) - Example "Me ... build-ing ... chairs, no, no cab-in-ets. One, saw ... then, cutting wood ... working ..." Wernicke’s aphasia - Also called posterior aphasia, receptive aphasia - Damage of Wernicke’s area - Mainly deficits in speech comprehension - Also production deficits related to semantics (speak fluent but nonsensical) - Example Examiner: What kind of work have you done? W. Patient: We, the kids, all of us, and I, we were working for a long time in the ... you know ... it's the kind of space, I mean place rear to the spedawn ... Conduction aphasia Damage of arcuate fasciculus Disconnection syndrome Mainly production deficits Problems producing and repeating speech Key Points Anatomy of Language: Anomia is the inability to find the words to label things in the world. It is not a deficit of knowledge. - A left-hemisphere network involving the frontal, parietal, and temporal lobes is especially critical for language pro- duction and comprehension. - The right hemisphere does have roles in language, especially in processing the prosody of language. - Language disorders, generally called aphasia, can include deficits in comprehension or production of language resulting from neurological damage. - Patients with Broca’s aphasia have problems with speech production, syntax, and grammar, but otherwise comprehend what is said or written fairly well. - The lesions that produce Broca’s aphasia may not be limited to the classically defined Broca’s area in the left inferior frontal cortex. - People with Wernicke’s aphasia have severe comprehension deficits but can produce relatively fluid speech; it is, however, rather meaningless. Originally linked to damage solely in Wernicke’s area (the posterior superior temporal gyrus), today Wernicke’s aphasia is also linked to dam- age outside the classic Wernicke’s area. - Aphasia can also result from damage to the connection between Wernicke’s and Broca’s areas (the arcuate fasciculus). Conduction aphasia is the disorder that results from such damage, and people with this type of aphasia have problems producing spontaneous speech as well as repeating speech. Language Comprehension Semantics Denotation Explicit or direct meaning Connotation Associated or secondary meaning Can be something suggested or implied by a word or thing Syntax - Refers to the way in which words in a particular language are organized into grammatically permitted sentences Phrase structure grammar Is a type of generative grammar in which constituent structures are represented By phrase structures rule or rewrite rule Chomsky’s transformational grammar Part of the theory of generative grammar Especially of natural languages Considers grammar to be a system of rules that generate exactly those combinations of words that from grammatical sentences in a given language Perceptual Analysis In order to understand spoken words, the listener has to decode the acoustic input Result of this acoustic analysis is translated into a phonological code Because through phonological code is how lexical representations of auditory words forms are stored in the mental lexicon After phonological format – lexical representations in the mental lexicon that match the auditory input can be accessed The best match can be selected Selected word – includes grammatical and semantic information stored in the mental lexicon Helps to specify how the word can be used in the given language Spoken word recognition processing proceeds anteriorly in the superior temporal gyrus (STG) Phenome processing appears localized to the left mid-STG Integration of phonemes into words appears to the left anterior STG Processing short phrases appears in the most anterior locations of superior temporal sulcus (STS) - - - Invariance Problem Refers to the challenge that listeners face when confronted with acoustic variability in speech sounds As they attempt to map these sounds to few phonological categories Coarticulation & segmentation problem Problem of text segmentation that occurs in some languages which are (traditionally) written without inter-word spaces Like Chinese and Japanese Compared to writing system which indicate speech segmentation between words by a word divider (space) Reading - Words can be symbolized in writing in three different ways: alphabetic, syllabic, and logographic Involves the visual analysis of horizontal lines, vertical lines, closed curves, open curves, intersections, and other elementary shapes Pattern recognition Selfridge’s pandemonium model Consists solely of stimulus-driven (bottom-up) processing and does not allow for feedback (top-down) processing Mclelland & Rumelhart’s connectionist model Proposed a computational model that has been important for visual letter recognition Assumes three levels of representation: A layer for the features of the letters of words A layer for letters A layer for the representation of words Permits top-down information (i.e., information from the higher cognitive levels, such as the word layer) Influence earlier processes that happen at lower levels of representation (the letter layer and/or the feature layer) Lexical Analysis Lexical access Refers to the stages of processing in which the output of perceptual analysis Activates word-form representations in the mental lexicon Including their semantic and syntactic attributes Lexical selection Is the next stage where the lexical representation in the mental lexicon that best matches the input can be identified (selected) Lexical integration Integrates words into the full sentence, discourse, or larger context Grammar and syntax are the rules by which lexical items are organized in a particular language To produce the intended meaning we begin by considering the mental lexicon, the brains store of words and concepts and ask how it might be organized and how it might be represented in the brain Speech Production Slips of the tongue Slips of the tongue help us to make models of language Anticipation Bake my bike Perseveration Cold curkey Substitution He is going up town Reversal/Spoonerism The lord is shoving leopard Levelt’s model Predicts the following result when someone is presented with a picture of a flock of goats and is asked to name them Conceptual preparation First the concept that represents a goat is activated Concepts related to the meaning of goat are also activated Lexical encoding Lexical selection Rules governing which lexical items may appear in which sentence structures Morphological encoding Contains both phonological information and metrical information Metrical information – information about the number of syllables in the word and the stress pattern Phonological encoding Ensures that the phonological information is mapped onto the metrical information Articulation Involves the posterior parts of Brocas area (BA44) Studies showed bilateral activation of motor cortex, the supplementary motor area (SMA) and the insula Can computers understand language? Turing test Alan Turing Test of a machines ability to exhibit intelligent behavior equivalent to that of a human ELIZA An early natural language processing computer program Created to demonstrate the superficiality of communication between humans and machines Simulated conversation by using a “pattern matching” and substitution methodology that gave users an illusion of understanding on the part the program Had no built-in framework for contextualizing events WATSON Question answering computer system capable of answering questions Combines artificial intelligence and sophisticated analytical software for optimal performance as a “question answering” machine Deductive Reasoning Conclusions follow certainly from the premises Premises A statement concerning only facts Do not have to be true, but rather empirical “some students pass this course” Conclusion “some students study for this course” Quantifier A word or statement concerning the premise that give it specific content “Some student study for this course” Atmosphere effect The atmosphere of a statement, or the context in which it is given Can influence the user’s ability to reason “All students drink alcohol, therefore all students are alcoholics” Categorical reasoning Consist of a premise followed by a quantifier Conditional reasoning Contains a premise and conclusion Syllogistic reasoning The most common form of deductive reasoning Consists of premises followed by a conclusion Modus ponens (Method of affirming) If P2 = p, then the conclusion = p Modus Tollens (Method of denying) If P2 = !P, then the conclusion = !P Denying the antecedent error P => Q: if not P, then not Q This is incorrect Affirming the consequent error P => Q: Q is true , therefore Q is true This is incorrect Wason selection task A common logical task Example Four cards are on the table, with a letter on one side and a number on the other. Tell the participants an arbitrary rule, and ask them which card they must flip over to test if the rule is correct This is a difficult task when implemented in a formal manner, but when it becomes practical, the success rate skyrockets to about 75% Inductive Reasoning From a large body of evidence, we make conclusions that are probably true Based on observation, yet the hypothesis cannot be officially proven (must use statistics to prove the probability of being true) Heuristics & Bias Heuristics are “rules of thumb” Availability heuristic We are biased by our experience with the availability of experiences, by recentness and frequency Representativeness heuristic We are biased by the way information is presented to us Even if the information is obviously correct/incorrect The manner in which it is presented is critical in our reasoning techniques Expertise Three stages of skill acquisition: Cognitive stage The processes of developing declarative encoding Associative stage Errors eliminated and good actions strengthened Autonomous stage No more thinking required Automatic actions Power law of learning 𝑇 = 𝑎𝑃 %& T = Time needed to solve a task A = Learning constant P = Practice b = learning constant Tactical learning Learning the sequence to solve a particular problem or perform a task Strategic learning Learning to organize the solution differently Deliberate practice Must be motivated to learn not just reproduce Given feedback on performance Focus on improving performance Intelligence IQ test - Began as a way to monitor childhood development Compared to an average Relative/Normative Relative Relating mental age to chronological age - - Subtests Factors - - Normative Relating scores to the average score Modern IQ tests are normative Spatial, verbal, non-verbal, reasoning, short/long term memory tasks Only one: g (IQ) Spearman’s model Multiple factors Thurstone’s Model (3-5) Guilford (120+) Crystallized & Fluid intelligence Fluid Intelligence: ability to solve new problems and identify patterns Crystallized intelligence: ability to use learned knowledge and experience Flynn effect! Every two years the general IQ goes up by 1 point Construct validity The debate over whether the structure or “construct” of the IQ test adequately tests true intelligence Or if there is some form of bias in this form of evaluation IQ tests biased towards who are not blind etc. Artificial Intelligence: Consistent A formal system is consistent if it does not contain a contradiction P and not P are both true Complete - A formal system is complete if all the statements can be made in a system are “decidable” or evaluable within that system - An artificially intelligent system Developed by Medin, Wattenmaker, and Michalski Was Heuristic (rule based search) using symbolic descriptions Medin, Wattenmaker & Michalski (1987) Categorization algorithm An algorithm that categorizes decisions and their outcomes Then decides its next move based upon the most rewarding outcome Category validity Humans tend to category validity Cue validity Cue Validity determines the value of a certain input/decision Based upon its probability of occurring or existing within a certain category Based on its particular feature, or cue INDUCE tends to cue validity Product- Process- equivalence DEEP BLUE Another artificially intelligent agent devised by IBM designed to play Chess Deep Blue beat the reigning chess grand master in 1997 The person it beat accused IBM of cheating, which they denied, fucked off, and retired Deep Blue Dijkstra Asking a computer whether it can think is like asking if submarines can swim INDUCE - - - Consciousness Core consciousness Brainstem Is the “star player” in the brain Houses some of the most essential brain functions for life Controls breathing, automatic responses in the face, neck and torso Is the channel between the brain and the PNS Thalamus RAS RAS = Reticular activating system Modulates arousal and attention through direct connections to the cortex Extended consciousness Cerebral cortex Sense of self, memories and expectations Clinical perspective - - Sleep What is consciousness? We are able to react purposefully to external stimuli and are awake Measured on a quantitative level Impairments Somnolence Abnormal sleepiness Acoustically arousable Sopor No sponatous movements Reaction to pain stimuli adequate Coma No reaction to any sort of stimuli Pain, sound, visual – nothing You can recover (Brain death) The body may be alive CNS (Central Nervous system) is unresponsive Vegetative state Includes failure to produce any purposeful voluntary behavior Unconscious, reflexive responses, opening eyes Unresponsive to stimuli (except pain) Abnormal sleep/wake cycles “normal behavior may persist Grinding teeth Smacking lips Transient: temporary Persistent: continuous Locked-in syndrome Patient is actually fully conscious Cannot move or make themselves clear to their environment Can make vertical eye movement or close eyelids Use EEG or Real Time fmri to communicate Terri Schiavo case Car accident Left in persistent vegetative state Fought in court over taking her off life support Was taken off and passed away Arguments were about whether she would ever recover Stages - Sleep comes in stages and fluctuates between different states Brain activity can be monitored via EEG (Electroencephalography) and EOG (Electrooculography) Stage 1 - Light sleep Where the person drifts in and out of consciousness Can be awakened easily Prepares the body for deeper stages pf sleep Stage 2 - Eye movements stop Brain activity lowers except for very rapid spikes in brain activity Sleep spindles Rapid spikes or spindles (visible on an EEG) During sleep stage 2 Generated in the Thalamus It is believed that is an automatic “test” the brain performs, trying to determine certain functions of certain brain structures when asleep K-complexes Singular, large spike in brain waves (visible on an EEG) Largest healthy spike in brain activity It is believed that these perform the function of initiating/performing memory consolidation Stages 3&4 This is deep sleep “Deeper than those conversations you have at 4AM after partying all night” Normal brain activity is extremely low, expect the brain emits strong Delta Waves Slow wave sleep Where the brain exclusively produces delta waves These are slow, large brain waves that are believed to be signs of neural development and reconsolidation/orientation These waves are strongest in people under the age of 40 - - REM/ paradoxical sleep REM = Rapid Eye Movement Clear sign of deep sleep The period when the most vivid dreams occur Increased activity in the Pons/Reticular formation Temporal and parietal cortex Decreased activity in the Visual, motor and prefrontal cortex Muscles are immobilized Dreaming “what you are doing right now when you imagine being at the beach instead of school” Eye movements and EEG The eye and its movements heavily affect EEG readings Can lead to distortion Allows to detect Rapid Eye Movement in sleep Sleep cycles through the stages Cycle change through the night We drift in and out of sleep cycles throughout the night Reaching stage 4 early during sleep Slowly drifting Back down to 2 and 3 Stage 1 before waking up again Functions of sleep Save energy Restore the brain Muscle repair Cell repair Tissue growth Protein synthesis Release of many of the important hormones for growth Sleep allows the body to repair and replete several cellular components that are needed for physiological functions Memory Strengthens and weaken connections Consolidate memories obtained that day REM sleep functions Function not entirely understood Used for memory maintenance NN of REM: memory ‘cleanup’ Neural Network model of REM Echidna does not dream Echidna does not dream Has an especially large cortex Binding problem What is the problem We don’t understand how all of the input from our senses, our memories and all parts of the brain bind together To form one cohesive experience Clinical neuropsychological evidence for the binding problem Apperceptive agnosia Shapes Spatial grouping Achromatopsia Colors Feature integration Prosopagnosia Faces Part binding Semantic dementia Semantic Conceptual binding Simultanagnosia (Balint‘s syndrom) Location binding Akinetopsia Serial Event binding Feature integration theory Binding is achieved in temporal dimension Via synchronous oscillation of neurons Binding by neuronal synchronization Synchronous oscillation is rhythmic repetitive neural activity in the CNS Neurons are firing at repeated intervals Creating oscillations in brain activity Free will - What you think you have Readiness potential During a conscious task Your muscles are “ready” about 1000ms before you actually move them Libet’s experiment Determine the relationship between Readiness potential and the subjective experience of free will Turns out we might not have free will but rather the inhibition to stop automatic responses RP starts early W Will to move W-S Corrected “will to move” EMG Automated muscle activity Soon’s fMRI experiment Showing a stream of letters every 500ms the user can press a button whenever desired you must remember the letter displayed whenever the button was pressed Determined that users were only able to decode their decision AFTER they performed it Qualia - Individual, separate instances of subjective, conscious thought Chalmers: easy problem & hard problem A thought experiment posed by David Chalmers where the experience of conscious thought poses two types of problems, easy and hard problems Easy problems problems - represented or explained by some behavior, habit, or function, such as controlling behavior, deliberately paying attention, the ability to recognize things in our environment can be explained, regardless of how rigorously, by observance of natural phenomena Hard problems problems that have an inherent subjective aspect, such as “why are we able to contemplate our own existence?” these questions are not fundamentally observable and therefore “hard” to prove. Note: The Hard/Easy problem experiment has been discredited and considered out of date with modern psychological understanding. Nagel What it’s like to be a bat Speculation about what it’s like to be a bat Our experiences would be completely different Because they perceive “visually” via echolocation Inverted Spectrum Problem The hypothetical thought experiment where two people experience the “same” color But in reality, see it differently We both know what red is, but do we see red the same way Originally posed by John Locke Mary the color scientist - Thought experiment where a hypothetical scientist “Mary” experiences her whole life in a black and white room Studies the science of visual perception using only black and white shading She knows what shade of “grey” is red, and which is blue The experiment poses the question: “Will she learn anything new by leaving her room and experiencing actual color for the first time?” Posed by Frank Jackson
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