Learning and Memory The behavior of organisms can be separated into three major categories: Reflexes Instincts Learned behaviors Reflexes - are involuntary responses to stimuli - have the advantage of producing rapid, reliable responses, but their inflexibility can be a disadvantage when the environment changes. Instincts - are automatic, but their resulting behaviors are more complex. - instinctive behaviors are consistent enough to be referred to as fixed action patterns. Learning - learning A relatively permanent change in behavior or the capacity for changes in behavior due to experience. - provides organisms with the most flexible means for responding to the environment. Types of Learning Repeated touching can result to habituation due to the reduced activity/excitatory potential at the synapse between the sensory and motor neurons in habituation was a direct result of the release of less neurotransmitter Habituation can last up to 3 weeks After Aplysia is sensitized by the administration of an electric shock to the head or tail, touching the siphon results in an enhanced gill-withdrawal response. Longer action potentials leads to the stronger gillwithdrawal reflex that we observe in sensitization. Aplysia Classical Conditioning-----ask (a) Touching the siphon (CS−) is not paired with shock and serves as a control. (b) Touching the mantle (CS+) is always followed by shock to the tail (UCS). (c) After several pairings of touching the mantle (CS+) followed by shock (UCS), touching the mantle alone now triggers gill withdrawal (CR). In the circled area in (b), the mantle shelf sensory neurons are sequentially activated, first by touching the mantle shelf and then by input from interneurons serving the tail. This sequence produces increased presynaptic facilitation, leading to the recording of greater postsynaptic potentials in the motor neuron than those recorded prior to training. 1. Associative Learning - occurs when an organism forms a connection between two features of its environment. example: Classical Conditioning 2. Nonassociative learning - A type of learning that involves a change in the magnitude of responses to stimuli rather than the formation of connections between elements or events. example: Habituation and Sensitization Habituation - occurs when an organism reduces its response to unchanging, harmless stimuli. Sensitization - occurs when repeated exposure to a strong stimulus increases response to other environmental stimuli. example: following major disasters such as earthquakes, people often experience exaggerated responses to movement, light, and noise. Classical Conditioning - A type of associative learning in which a neutral stimulus acquires the ability to signal the occurrence of a second, biologically significant event. Conditioned Stimulus (CS) refers to an environmental event whose significance is learned, whereas an (learning; bell sound) Unconditioned Stimulus (UCS) has innate meaning to the organism. (unlearned; food) Conditioned responses (CR) are those behaviors that must be learned (salivating to bell sound) Unconditioned Responses (UCR) appear without prior experience with a stimulus. (salivating to food) Using Invertebrates to Study Learning sea slug = Aplysia californica Brains = Neural net Touching the animal’s siphon reliably produces a protective response known as the gill-withdrawal reflex, in which the gill is retracted. Classical Conditioning in Vertebrates fear responses in rats and eyeblink conditioning in rabbits. Classical Conditioning of Fear Amygdala plays a role in classical conditioning of emotional response such as fear. 1. Rat hears a tone (CS) 2. Electric shock (UCS) 3. Stops eating and shows fear 4. Association is made between the tone and the shock 5. When the rat hears the tone the amygdala sends out a strong response for fear in the absence of shock Classical Conditioning of the Eyeblink 1. Rabbit hears a tone (CS) 2. Followed by a puff of air in the rabbit’s eye (UCS) causing movement in the nictitating membrane. Nictitating Membrane - An additional, moveable inner eyelid found in some birds, fish, and mammals but not in humans. 3. After several pairing of tone and puff the rabbit blinks when they hear the tone even without the puff of air. the importance of the interpositus nucleus in classical conditioning shows how during learning there is an increase in the structure’s response to tone/CS. Inactivation of this structure can inhibit classical conditioning Interpositus Nucleus - cerebellar nucleus thought to be essential to classical conditioning in vertebrates. Lesions can change our behavior due to loss of that structure and this change can be due to the lack of info travelling via fibers affected by the lesion. Interpositus nucleus is primarily responsible for the formation of the classically conditioned response of the nictitating membrane in rabbits; red nucleus only suppresses but doesn’t prevent learning. Cerebellum damage and shrinkage is correlated with speed of acquisition of CR Cerebellar Circuits and Classical Conditioning Purkinje cell - A cell in the cerebellum that influences its activity by forming inhibitory synapses with the output cells in the deep cerebellar nuclei. - receives input from climbing and parallel fiber Climbing Fiber - A fiber originating in the inferior olive of the brainstem that forms synapses on the large Purkinje cells of the cerebellar cortex. Parallel Fiber - A fiber originating in the granule cells of the cerebellum that synapses on the Purkinje cells. - originate from granule cells Granule Cell - A cell within the cerebellum that is the source of parallel fibers. - receive input from mossy fibers Mossy Fibers - integrate info from parallel and climbing fiber. - Originate from neurons in the pons. Purkinje cells form inhibitory synapses, located in the deep cerebellar nuclei therefore influences the output of the cerebellum learning will occur if the climbingfiber and parallel-fiber synapses onto a Purkinje cell are activated at the same time. Paired stimulation resulted in reduced activity in Purkinje cell EPSPs. The reduced activity in the Purkinje cells is known as long-term depression, or LTD. 1. Three Events Occur: When the climbing and parallel fibers are active, three things happen simultaneously in the Purkinje cell: calcium (Ca2+) and sodium (Na+) ions enter the cell, and a chemical known as protein kinase C (PKC) is activated. 2. Fewer Glutamate Receptors: These events together lead to a decrease in the number of glutamate receptors in the Purkinje cell membrane. Since there are fewer receptors to respond to glutamate, the result is a reduction in EPSPs. 3. Connection to Learning: Even though a reduction in activity might seem like it wouldn't help with learning, this process of long-term depression (LTD) is actually crucial for certain types of learning, like the eyeblink conditioned response (CR), which involves the inferior olive and climbing fibers. This reduction in EPSPs due to fewer receptors is part of how this learning happens. Trace Conditioning and Extinction Delay Conditioning: The CS and UCS overlap; the CS is continuous until the UCS is presented. This type of conditioning is generally easier to learn because the stimuli are presented closely together. Example: The dogs learn to associate the tone with the food, causing them to start salivating at the sound of the tone, even before the food is presented. Trace Conditioning: There is a gap between the end of the CS and the beginning of the UCS. This requires the learner to remember the CS during the gap, making it generally harder to learn but offering insights into the learner's memory and cognitive abilities. Example: The dog will start salivating when they hear the bell, even though it has stopped before the food arrives. Trace conditioning requires the forebrain structures to hold the CS information during the stimulus-free interval before transmission to cerebellum NOTE: Classical conditioning in vertebrates involves the amygdala (emotional learning) and the cerebellum (skeletal reflexes). - In addition to the cerebellum, trace conditioning requires activity in the forebrain. Also require conscious, declarative processes Memory Information Processing Models - theories of memory that seek to explain the management of information by the brain, from detection to storage to retrieval. 3 types of Long-Term Memory 1. Semantic memory - contains basic knowledge of facts and language. Ex. “who is your mother?” 2. Episodic memory -relates to your own personal experience. You use your episodic memory to remember the episodes of your life. Ex. “where were u last Monday?” 3. Procedural memory - stores information about motor skills and procedures. Ex. riding a bicycle, using a software program Semantic and episodic memory are both “Declarative Memories” because they can easily be described with words/declared. Procedural memories are difficult to verbalize but easy to demonstrate or perform Declarative memories: conscious and explicit recall involved Procedural Memories: unconscious and implicit recall involved Classical conditioning, habituation, and sensitization are also considered examples of nondeclarative processes. People with anterograde amnesia suffer from explicit memory impairment but have an intact implicit memory Engram - A physical memory trace in the brain. Brain Mechanisms in Memory Lashley posed that no single area is more responsible for learning and memory than any other. Penfield’s study shows possible role for the temporal lobe in the formation and retention of long-term memories. conclude that hippocampal spatial maps are formed within minutes of entering a new environment In a study where they deleted he CA1 gene that has a role in the NMDA receptors has a negative impact on LTP Disrupting “second messengers” transmission of signal also negatively impacts LTP When LTP is impaired and is in a same place, a new map is formed instead of reactivation of old one Spatial learning can also be impaired if a NMDA antagonist is administered The Diencephalon and Memory Composed of the thalamus and hypothalamus Patient N.A. suffered ffrom antero and retrograde amnesia after being stabbed by a foil fencing sword in the nose. Short term memory intact, long term impaired. Korsakoff ’s syndrome - Anterograde amnesia resulting from thiamine deficiency, typically found in chronic alcoholics. Semantic Memory and the Cerebral Cortex “convergence zone,” or a particular location responsible for assembling separate aspects of a memory into a whole. convergence zone for semantic memories are areas along the left lateral inferior frontal gyrus. Most active when incorrect language/worl knowledge was used The Temporal Lobe and Memory Patient H.M. proved that memories do not reside in the medial temporal lobe itself necessarily essential for the retrieval of stored memories. Delayed Nonmatching To Sample (DNMS) Task - A standard test of memory in which the subject must identify the novel/new member of a stimulus pair following a delay. (Monkey example) Study showed that lesions does not affect short term memory but shows difficulty in forming long term memories Long-Term Potentiation (LTP) Long-Term Potentiation (LTP) is a process where repeated stimulation strengthens the connections between neurons, leading to more efficient signal transmission. Example: Playing and practicing a new note Ammon’s horn is further divided into four sections, named CA1 to CA4. (CA stands for the Latin term for Ammon’s horn, cornu Ammonis.) Hebbian synapse and LTP require relatively simultaneous firing called: Associativity A condition believed necessary for learning in which the pre- and postsynaptic neurons are nearly simultaneously active. Cooperativity A condition for the formation of LTP in which several synapses onto the target postsynaptic neuron must be simultaneously active. Both are needed for glutamate to influence postsynaptic neuron LTP and Spatial Memory Spatial memory, or an organism’s ability to map a location Episodic Memory and the Cerebral Cortex Patients with a damaged prefrontal areas of cortex suffer from source amnesia. Source Amnesia - memory loss for the circumstances in which a particular fact or skill was learned. Anterior prefrontal cortex and the Costerior cingulate cortex participate in retrieval of personal episodic memories. we consult our personal experience to determine reality We utilize episodic memories to distinguish between fantasy and reality When considering fantasy, ares semantic processing shows the greatest activity because of violation of world knowledge. Short-Term Memory and the Brain 4 components 1. Central Executive Attentional aspects in STM Neural basis are dorsolateral prefrontal cortex (DLPFC) and the anterior cingulate cortex (ACC) Lesions hinders the switching of attention to new things Example: Card sorting experiment 2. Phonological Loop 3. Visuospatial Scratchpad 4. Episodic Buffer level of maturity in the prefrontal cortex is necessary for short-term memory. And area of lesion (prefrontal) affects STM Large memory capacity = more activation of ACC; use imagery of story telling Less memory capacity = less activation of ACC; use repetition, rehearsal to remember The Striatum and Procedural Memory Straitum includes the basal ganglia (motor patterns) and nucleus accumbens (reward for learning new procedures) Rats with lesion in the hippocampus experienced lowered explicit memory but if the lesion is in the basal ganglia procedural memory is affected The Effects of Stress on Memory Freud believed that traumatic memories can become so inaccessible that they are essentially lost or repressed PTSD - traumatic memories are vivid and intrusive the Yerkes-Dodson Law indicates that there's a sweet spot for arousal levels that leads to optimal performance. Too little arousal might lead to lack of motivation or boredom, while too much can cause stress and hinder performance. The key is finding the right balance based on the task at hand. In this case, the amygdala LTP successfully processed the emotional component of the stressful event, but the hippocampus LTP (damaged by lack of oxygen) was unable to encode the declarative details of the event. Example: Man buried in the sand Stress affects the hippocampus, amygdala and the functions normally carried out by the prefrontal cortex, including coping skills, decision making, planning, and multitasking. chemicals known as glucocorticoids, including cortisol, are released during stress and increase the activity of the amygdala Higher levels of cortisol are correlated with the reporting of more false memories Propanolol, which blocks the effects of glucocorticoids in the brain, might prevent the formation of traumatic memories when administered immediately following a traumatic event Age & Memory As people age, decreased blood flow is observed in many parts of the brain essential to memory and cognition, especially in the frontal and temporal lobes. However, the aging brain also shows areas of increased activation. suggest that these areas of increased activity represent a reorganization of the brain that allows cognitive performance to remain stable in spite of age-related deficits in brain function