The document discusses the history and current state of research on the physiological components of learning and memory. It notes that while early research knew little about these topics, recent decades have seen many advances, though much remains unknown. The document goes on to describe various processes involved in memory formation and storage at the molecular, cellular, and neural network levels. It also discusses theories of memory like Hebbian plasticity and distinguishes between types of learning like classical and operant conditioning.
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The document discusses the history and current state of research on the physiological components of learning and memory. It notes that while early research knew little about these topics, recent decades have seen many advances, though much remains unknown. The document goes on to describe various processes involved in memory formation and storage at the molecular, cellular, and neural network levels. It also discusses theories of memory like Hebbian plasticity and distinguishes between types of learning like classical and operant conditioning.
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65 years ago, a great scientist, E.R.
Hilgrad, said: "This is a negative aspect
of science that after many years of research we know little about the physiological components of learning." It's been more than half a century since those words were written. There are 65 years in which research has been directed towards the discovery of "physiological components of learning." Surprisingly, we know very little about what is happening in the brain, about how learning and memory are carried out. However, in recent years have appeared many firsts. The horizon begins to be brighter. Perhaps, in the first decade of the XXI century we started really to hope that we will find a solution in this domain. Furthermore, the hope comes from studies in molecular and biophysical level. The solution, in terms of memory, is based on a high capacity for synthesis of knowledge in the field of neurobiology. This involves the flow of ions, secondary messengers, gene suppression, channel proteins, membrane structure. Are also involved levels beyond the molecular structure and ultrastructural. In mammals, the complex anatomy of the brain is heavily involved. Are also involved neural networks of interest for thousands if not millions of neurons, millions if not billions of synapses. Memory is a process by which humans and other organisms encode, store and reuse information. Memory is a psychological process of reflection of past experience by setting (imprint and retention), recognition and reproduction of sensory images, ideas, emotional states or movements of the past. Memory processes are conducted both in relation to sensory reflection and in relation to thought processes and language, to some people predominating sensory-intuitive memory to others the verbal-abstract. Recognition and reproductionas memory processes are conditioned by the attachment process, of memorizing. Memorization can be intentional (voluntary) or unintentional (involuntary). Recognition is performed in the presence of sensory or verbal stimuli which have acted before, while reproduction is achieved without these stimuli. The inverse phenomenon of memorizing is forgetting, which is manifested by the inability to reproduce or to recognize, and sometimes by incorrect reproduction or recognition. Can be differentiated logical and mechanical memorizing, but specific for a person is logical memorizing, based on understanding of the saved. Memory and learning terms are difficult to delimit, as they have many common features. Let’s try in the following to specify the areas of the two notions. We saw above that memory is the psychological process consisting in the retention, preservation, reproduction of facts from our previous experience. Let's see the features of learning: A) While memory is primarily a way of reflecting, learning is defined notably as an activity through which we fix knowledge in memory. B) Learning is something more complex than simple memorization. Learning involves an effort of understanding, systematization and organization of knowledge, exercises of their use in practice. C) Learning covers a wider range of phenomena than simple memorization. For example, a student solving mathematics problems, he perfects, through these, the ability to solve problems, to use the mathematical knowledge in an independent and original way. As results from the above, the main processes of memory are: imprint, the maintenance and updating. We will deal on all of these processes. a) Memorizing - the first process of memory, in which occurs the imprint and fixing of our experience. In the process of memorazing, the nerve connections systematizes and constitutes in dynamic stereotypes. As you remember, in a dynamic stereotype is about a series of documents integrated into a meaningful whole unit. The words of a poem is not strung, simply out our memory, as a string strung beads. b) Retention - once memorized the facts are kept a shorter or longer time until they are replicated or serve in an act of recognition. Keeping it so the second process of memory and is in the range of store and update stored. During storage, our knowledge go through a manufacturing process and systematization. c) Update - is the third process of memory, a process in which those stored in our memory some time are taken out and used, recovered. Updating can be done by recognition or reproduction. Encountering an object, a fact in the past, we recognize it, ie we realize that we had to deal with it in our previous experience. Recognition may be vague or precise. We sometimes encounter people who appear as familiar faces. But we only have this feeling of something known, familiar, but we can recount something about that person (who's, where I met etc). It is vague or generic recognition. Sometimes recognition is more accurate. Memory is found as: imaginative memory - by preservation, reproduction of the past images; verbal memory - logical - by preserving ideas, information; most controversial is emotional memory - remembering past emotions; Memory can be divided conventionally into three subcategories: sensory memory, motor and central. Sensory memory is for instance the ability of visual cortical areas to assemble and retain spatial patterns for further memory. Motor memory is the brain's ability to learn and retain information for subsequent execution of some actions. Main memory is the brain's ability to assemble, integrate and make sense of sensory information and store them for future use if needed. An important feature of these three types of memory is the brain's ability to make associations. A graphic demonstration of the association was given by Pavlov, who trained a dog to associate the sound of a bell with food, the has dog drooled upon hearing the bell. Here, the brain made association of various sensory information with food. During this conditioned process, the second way (S2) becomes operational, probably due to the activity of association areas of the brain. Thus, concomitant repetition of two presynaptic systems reinforces the auditory path, a phenomenon predicted by Hebb 50 years ago. Proof of Hebb's synapse was made by the NMDA receptor discovery. As presented in the previous chapters, NMDA receptor binds to central excitatory neurotransmitter glutamate, and is both a channel voltage dependent and ligand-dependent one. In order to be open postsynaptic cell must be depolarized, and the receptor must link the glutamate. Depolarization of these synapses is due to glutamate release from presynaptic sites, this also binding to other receptors on the postsynaptic surfaceso called kainate receptor, a receptor for glutamate in the CNS frequently (Fig. 8-3). Hebb’s synapses were found in the dendritic thorns of neurons in the hippocampus, pyramidal and stellate cells and Purkinje cells of the cerebellum. If the opposite terminal of the NMDA receptor releases glutamate, it binds to NMDA receptor, but does not open the channel which is blocked by Mg2 + ions. But if the terminal found near the kainate receptor releases glutamate, postsynaptic membrane is depolarises. Therefore the lock of channel carried out by Mg2 + ions is removed and the channel opens, allowing Ca2 + ions to penetrate into the cell. The presence of intracellular Ca2 + activates protein kinases from postsynaptic cell, leading to the generation of nitric oxide (NO), which diffuse into the presynaptic cell and induces even more the release glutamate. Alternatively, NO can lead to the synthesis of several kainate receptors. The process was called long-term potentiation (long time potentiation - LTP). Learning is a process in which the brain accumulates information from the external environment. This definition excludes instinctive knowledge (intrinsic), with which we were born. Learning was divided into associative and nonassociative. Nonassociative learning was further divided into skill and awareness. The skill is the attenuation of a behavioral response to repeated application of non-toxic stimuli. For example, we flinch when the door is slammed, but not going to flinch when the door is closed several times during a test. Sensitization is an increased response to a noxious stimulus. For example, touching the skin gently that has previously been subjected to a painful stimulus will lead also to a painful response. Associative learning - is divided into classical conditioning (Pavlovian) and operant conditioning (instrumental). The concept was first introduced by Pavlov which created the association of two stimuli in the mind of a dog - one conditioned and the other unconditioned. Conditioned stimulus is an applied stimulus that can be auditory (eg a voice or a bell) or visual (a light). Unconditioned stimulus is a stimulus that has not been tested by
an experimenter. To the observation food (unconditioned stimulus),
the dog drools in a reflexed way (unconditioned response). The answer is unconditional because it is not innate and not
acquired through learning.
An association between the two stimuli is achieved in the dog
mind if the conditioned stimulus (light) is followed repeatedly by
unconditioned stimulus (food sightings). Operant conditioning (instrumental) - is an important form of associative learning. The animal learns that by a certain behavior manages to get a favorable response (receive a reward), or it will be punished. For instance if the reward is pleasant - food - then the animal will repeat its action. If the reward is unfavorable, it will not repeat the action, it avoid. Operant conditioning (stimulus-response learning) allows the enrichment of precise motor ways of reaction to certain stimuli. Stimulus-response learning can create a new reaction by combining "step by step" of some preexisting elements in the stock of knowledge of the subject, or selective reinforcement of spontaneous manifestations occurring in its behavior. For example, step by step learning is writing a letter as its made in school, learning a typical ballet positions (when the teacher instructor actually sits with his hand the subject feet in the standard position), learning to adjust the microscope (when the teacher is leading the hand of the subject in matching the inclination of objective or positioning the preparation so that student to obtain the optimal clarity), effective use of instruments or tools, etc.. Internally there must be an terminal act that causes satisfaction to the learner, an effect that strengthens the internalized connection. Terminal act may be an innate one (satisfying the hunger) or one acquired (praise). Feeling satisfaction is to do something with pleasure or obtain a reward (food, a desired object, praise, weakening of pain). This condition expresses the law of effect, which says that "when it forms a modifiable connection between stimulus and reaction and this is followed by satisfaction, the link is strengthened." Strengthening a stimulus-response connections is indicated by increased frequency of a particular response to a particular stimulus or the diminution of an previous tendency. Externally - accelerating learning occurs by reducing the time between the emergence of learned response and the appearance of strengthening. Burrhus Frederich Skinner described one of the most influential systems for studying the learning process. He was interested in explanations by contingency, "which leads to which" - and therefore he deals with the anticipation of behavior more than its understanding, result so that his sistem is not a theory in the strict sense of the word. During the researches, he developed highly efficient methods for controlling behavior, and therefore he handles with the training methods. His ideas are of general interest to those interested in educational processes that have led to special methods of training, such as learning machines and programmed learning. However the validity of his method known as "operant conditioning" appears most clearly in experiments on animals. He started from the reflex conditioning of Pavlov's - the process of forming a new reflex or temporary connections by associating repeated of a unconditioned stimulus (bell sound) with indifferent stimulus (food). Skinner, respecting the same conditions, revealed a new kind of conditioning, different from pavlovian one, which he calls "operant conditioning." Operant conditioning significance consists in the fact that it shows a second way of handling the curing process (of pressure, drawing, spinning), and thus the subject get a reward. Is famous the "Skinner's box" (fig.8-5) in which a rat receives food if press a lever that opens the box. In these experiments, the animal must issue an appropriate response in order to produce the strengthening stimulus, and thus to receive the food. Unlike the classical method, hungry rat must press the lever, if not done so, the food is not presented. The animal investigates free and during this activity issues a response that changes the environment so that its interior behavior is affected. These changes in the environment could be called rewards, but it is usually preferred the term "strengthening stimulus." When such stimuli follows an answer, they increase the likelihood that the animal to behave in the same way again. Thus, the animal learns and its behavior is modeled in a way that is more efficient in the operation of environment. Sometimes one part "irrelevant" of the behavior occurs before the response that provides reward. In 1948 Skinner calls it "suspicious behavior" because the monkey behave as is obliged to do this to get the reward even though it was not planned such juxtaposition of answers in advance. Skinner's experimental analysis, of the behavior of animals, has reached a stage that allows the study of complex systems in lower organisms at a higher level of scientific rigor. Memory can be a "labile" process short-term memory (STM) and a "stable" process long-term memory (LTM). Short-term memory lasts for seconds, minutes or hours. It's the kind of memory that we use when looking for a phone number to dial, and the number will not remain in memory longer than a few minutes. If we retain all phone numbers that we have formed in a lifetime we might not to be able to focus on important issues. The transiency and lability of short-term memory suggests that it has a "physiological" base - perhaps a transient alteration of the synaptic "resistance" of certain pathways in the brain. Long-term memory lasts for days, years, decades. This holds important information about the life of an animal - where to find food or a mate, how to avoid a predator or poison. It is believed that in general the information to be stored first passes through STM and then in LTM. This transfer is called consolidation. We have already noted that STM can be distinguished from LTM by its great "lability". It is not difficult to interrupt STM, in contrast, LTM is difficult to replace. It follows that we can prevent the consolidation by STM disorder. This can be done by exposing the brain to various "hazards": concussion, electroconvulsive shock (SEC), hypothermia, certain medications, etc. There are two forms of memory: declarative memory, explicit which stores the information that can be reused at any time consciously. It is needed to recognize certain things - animals, faces, fruit. Implicit memory, procedural does not require conscious activation for storing and recall. It is important for example to learn to play the piano. To form declarative memory, first information reaches the cortex association via primary sensory cortical area. From here, on the path area 28, the information reaches the hippocampus, which is important for long-term storage of declarative memory. By associating the diencephalon structures and prefrontal cortex information is again stored in the association cortex Information can be transferred from the STM to the LTM through repetition. Particularly even this transfer from the STM level to LTM is involved in injuries of the above structures which lead to the emergence of neurodegenerative diseases (eg Alzheimer's disease). In addition, memory storage can be blocked by means of an electric shock. The most important neurotransmitter in hippocampus is glutamate. Memory formation is induced by norepinephrine and acetylcholine. Damage the hippocampus or its connections lead to anterograde amnesia. Patients affected will remember previous things but not subsequent event. Korsakoff syndrome (frequently in the chronic alcoholism) can occur both anterograde and the retrograde amnesia. Procedural memory (implicit) is not associated with damage to the hippocampus. Depending on the task, are involved the basal ganglia, cerebellum, amygdala and cortical areas. Both the basal ganglia and the cerebellum plays an important role when we learn occupations. The afferent impulses reach the cerebellum through pontini and olivary nuclei. The storage capacity of the cerebellum can be lost for example through toxic injury, degenerative diseases and trauma. Dopaminergic projections of substantia nigra plays an important role in procedural memory formation. The amygdala is important for the conditioning of anxiety reactions. It receives information from the cortex and thalamus and influences motor and autonomous functions (eg. muscle tension, palpitations) through the reticular formation and hypothalamus. Bilateral removal of the amygdala with portions of the hippocampus and temporal lobe lead to amnesia and to uninhibited behavior (Kluver-Bucy sdr.).
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