Learning is not just the remembrance of some initial intense stimulation. It is even more importantly the remembering of associated experiences. (Eccles; Evolution of Brain, 150)

Learning and Memory. (Kandel; Principles of Neural Science, 1227)

Learning can lead to structural alterations in the brain. (Kandel; Principles of Neural Science, 1274)

Changes in the efficacy and local geometry of connectivity provide a basis for learning and memory. (Purves; Neuroscience, 599)

Learning feelings of like or dislike requires the amygdala; learning habits requires the neostriatum; learning a discrete motor response to a conditioned stimulus requires the cerebellum. (Squire & Kandel; Memory, 212)

Area10 in the lateral prefrontal cortex is involved with memory and planning, cognitive flexibility, abstract thinking, initiating appropriate behavior and inhibiting inappropriate behavior, learning rules, and picking out relevant information from what is perceived through the senses. (Gazzaniga; Human, 20)

Arts may be a useful form of learning.  It has been suggested that arts help us to categorize, increase our predictive power, react well in different situations. Thus, arts do contribute to survival. (Gazzaniga; Human, 226)


Research Study — Strengthening Synaptic Connections by NMDA for Long-Term Potentiation (LTP)

Research Study — Hippocampal CA3 network Pattern Completion

Research Study — Sleep promotes formation of Dendritic Spines

Research Study — Plasticity of Basket-Cell Network Regulates Adult Learning

Research Study — NMDA Receptor Ion Channel Crystal Structure

Research Study — Hippocampus Dendritic Inhibition Supports Fear Learning

Research study — Entorhinal–Hippocampal Ensemble Activity during Associative Learning

Research study — Dopamine Prediction Errors Circuitry


Synaptically connected neural patterns are the brain’s representation of ideas. These synaptically connected neural patterns are ingrained via extensive training and learning. The synaptically connected neural patterns are sparsely distributed throughout the brain. Individual neurons will be used and reused many times in different synaptically connected patterns. An example of this kind of connectivity has been found in the motor cortex.


Advantage of Learning

Advantage of learning; adapt to rapidly changing aspects of environment; could not be achieved by natural selection alone. (Johnston; Why We Feel, 81)

Phylogenetic learning - achieved through natural selection over generations;  Ontogenetic learning - arising from an individual's unique experiences. (Johnston; Why We Feel, 33)

Learning could complement biological evolution by allowing individuals to discover how to survive and reproduce within their own unique and changing environment. (Johnston; Why We Feel, 82)

Learning is a mechanism by which new events can evoke our inherent repertoire of positive or negative feelings, based on their association with the primary set. (Johnston; Why We Feel, 106)

Experience Causes Synapse Patterns to Strengthen

Each and every new experience causes the neuronal firing across some synapses to strengthen and others to weaken.  The pattern of change represents an initial memory of the experience. (Ratey; User's Guide to Brain, 191)

Long-term potentiation (LTP) is the cellular mechanism that causes synapses to strengthen their connections to one another, coding an event, stimulus, or idea as a pattern of connections. (Ratey; User's Guide to Brain, 191)

LTP blazes a new trail along a pattern of neurons, making it easier for subsequent messages to fire along the same pattern  The more the pattern is refined, the more permanent the message -- the new learning -- becomes. (Ratey; User's Guide to Brain, 191)

As neurons in the chain strengthened their bonds with one another, they began to recruit neighboring neurons to join the effort.  Each time the activity is repeated, the bonds become a little stronger so that eventually an entire network develops that remembers the skill, the word, the episode, or the color.  At this stage, the subject becomes encoded as memory. (Ratey; User's Guide to Brain, 191)


Research study — Dendritic Spines and Memoriesin the mouse cortex, learning and novel sensory experience lead to spine formation and elimination by a protracted process.


Inhibition Prevents Distracting Noise

Inhibition is a crucial part of learning because it prevents noise from distracting our focus on what matters. (Ratey; User's Guide to Brain, 196)

Forgetting Removes Trivial Memories

Although forgetting can be frustrating, it is a necessity -- otherwise memories would clog our minds. (Ratey; User's Guide to Brain, 196)

Habituation, Sensitization, and Classical Conditioning

Three simple forms of learning -- habituation, sensitization, classical conditioning. (Eichenbaum; Neuroscience of Memory, 41)

Elementary forms of learning: habituation, sensitization, and classical conditioning. (Kandel; Principles of Neural Science, 1248)

Habituation -- simplest form of implicit learning, an animal learns about the properties of novel stimuli that are harmless. (Kandel; Principles of Neural Science, 1248)

Habituation - loss of arousal as a result of repeated exposure. (Johnston; Why We Feel, 104)

Synaptic depression of the connections made by sensory neurons, interneurons, or both is a common mechanism for hibituation. (Kandel; Principles of Neural Science, 1249)

Sensitization -- when an animal repeatedly encounters a harmful stimulus, it learns to respond more vigorously, even to similar harmless stimuli. (Kandel; Principles of Neural Science, 1250)

Sensitization is more complex than habituation. (Kandel; Principles of Neural Science, 1250)

Classical conditioning is a more complex form of learning than sensitization.  Rather than learning only about one stimulus, the organism learns to associate one type of stimulus with another. (Kandel; Principles of Neural Science, 1252)

Learning without Awareness

Active neurons in the cortical system that are apart from awareness activity at the moment can still lead to behavioral changes but without awareness. These neurons are responsible for the large class of phenomena that bypass awareness in normal subjects, such as automatic processes, priming, subliminal perception, learning without awareness, and others. (Koch, Neuronal Theories; Koch and Crick; Neuronal Basis, 97)

Procedural Memory or Skill Learning

Procedural memory or skill learning is the first memory function to develop in the infant brain's early stages of growth. (Ratey; User's Guide to Brain, 200)

Implicit memory is responsible for the laying down of skills and habits that, once learned, do not have to be consciously thought about, such as eating, talking, walking, riding a bike, and the way to go about making friends. (Ratey; User's Guide to Brain, 200)

Implicit memory provides stereotyped but extremely reliable  movements, and involves the basal ganglia and cerebellum. (Ratey; User's Guide to Brain, 200)

Motor Learning

Cellular mechanisms of Learning and a Biological Basis of Individuality. (Kandel; Principles of Neural Science, 1247)

Procedural memory - reflects motor learning. (Edelman; Wider than the Sky, 51)

An attractive hypothesis is that collaborations between the cortex and basal ganglia set up synaptic changes that lie behind procedural learning. (Edelman; Wider than the Sky, 93)

During conscious learning of tasks, a considerable amount of cerebral cortex is engaged. (Edelman; Wider than the Sky, 91)

Motor programs are refined continuously by learning. (Kandel; Principles of Neural Science, 672)

Brain's Motor Function and Learning

Brain's motor function affects so much more than just physical motion.  It is crucial to all of the brain functions -- perception, attention, emotion -- and so affects the highest cognitive processes of memory, thinking, and learning. (Ratey; User's Guide to Brain, 175)

Motor programs continually reorganize into sequences of motor movements that reflect what we learn each time, to lead to a well-thought-out and successful performance.  We are always modifying and learning through movement. (Ratey; User's Guide to Brain, 176)

Sequencing is a motor activity that involves manipulating and organizing the serial order of information and integrating this information with previously learned data. (Ratey; User's Guide to Brain, 177)

Movement Is a Fundamental Basis of Learning

Movement is a fundamental basis of learning. (Ratey; User's Guide to Brain, 204)

For motor memories, prefrontal cortex plans and organizes events while the basal ganglia and hippocampus act together to store the memories for the long-term. (Ratey; User's Guide to Brain, 205)

Motor memory and skill-learning are intimately interrelated. (Ratey; User's Guide to Brain, 205)

One reason motor function and memory are so closely linked, is that they are both coordinated by the frontal lobe. (Ratey; User's Guide to Brain, 206)

Physical movement helps cement the learning. (Ratey; User's Guide to Brain, 270)

Cerebellum in Learning

Purkinje cells of the cerebellum have long term depression of synaptic transmission (LTD), a form of synaptic plasticity that may underlie certain forms of motor learning. (Kandel; Principles of Neural Science, 240)

Cerebellum is important for maintaining posture and for coordinating head and eye movements, and is also involved in fine tuning the movements of muscles and in learning motor skills. (Kandel; Principles of Neural Science, 322)

Cerebellum circuits are involved with the timing and coordination of movements in progress and with the learning of motor skills. (Kandel; Principles of Neural Science, 665)

Synaptic transmission in the cerebellum's circuit modules can be modified, a feature crucial for a motor adaptation and learning. (Kandel; Principles of Neural Science, 832)

Damage to the cerebellum disrupts the spatial accuracy and temporal coordination of movement.  It also markedly impairs motor learning and certain cognitive functions. (Kandel; Principles of Neural Science, 833)

The kind of motor adaptation and learning with which the cerebellum is concerned requires trial-and-error practice.  Once the behavior becomes adapted as learned, it is performed automatically. (Kandel; Principles of Neural Science, 849)

Lateral cerebellum appears to be particularly important for learning both motor and cognitive tasks in which skilled responses are developed through repeated practice. (Kandel; Principles of Neural Science, 850)

Basal Ganglia in Learning

Neurons in the basal ganglia regulate movement and contribute to certain forms of cognition such as the learning of skills. (Kandel; Principles of Neural Science, 331)

Neurons in the basal ganglia regulate movement and contribute to certain forms of cognition such as the learning of skills. (Kandel; Principles of Neural Science, 331)

Brain stem integrates spinal reflexes into a variety of automated movements that control posture and locomotion. (Kandel; Principles of Neural Science, 672)

Several interconnected areas of cortex that project to the descending systems of the brainstem and spinal cord initiate and control complex voluntary movements. (Kandel; Principles of Neural Science, 672)

Orbitofrontal Cortex in Learning

Research study — Orbitofrontal  Learning Using Inferred Valuesorbitofrontal cortex is fundamental for accessing model-based representations of the environment when value is computed or inferred on the fly when it is needed.


Learning a New Skill

When people initially learn a task, the prefrontal cortex -- involved in short-term memory and many kinds of learning -- is relatively active. (Ratey; User's Guide to Brain, 179)

Research studies suggest that a newly learned skill can be impaired, confused, or even lost if a person tries to learn a different motor task during the critical 5 to 6 hour, when the brain is trying to stabilize the neural representation and retention of the original task. (Ratey; User's Guide to Brain, 179)

Movement and Learning

Extensive linkages between movement and learning. (Ratey; User's Guide to Brain, 180)

Gesturing and pantomime speed up the process of learning to talk. (Ratey; User's Guide to Brain, 180)

Babies make certain gestures before they can say the corresponding words. (Ratey; User's Guide to Brain, 180)

Children who signed were found to be significantly ahead of those who didn't in acquisition of vocabulary as well as in cognitive and IQ test. (Ratey; User's Guide to Brain, 180)

Signers showed high motivation to communicate and talked earlier than non-signers. (Ratey; User's Guide to Brain, 180)

Walking impacts human development as it facilitates cognition, spatial relations, communication, and social ability. (Ratey; User's Guide to Brain, 180)

Normal babies begin walking between the ages of nine and seventeen months. (Ratey; User's Guide to Brain, 180)

Nucleus Accumbens Important for Learning

Nucleus accumbens is important for learning, in part because it tags information with a signal of intensity that tells the rest of the brain to pay attention. (Ratey; User's Guide to Brain, 244)

Brain's reward center is central to learning and provides a motivation for doing something or the sense of feeling satisfied. (Ratey; User's Guide to Brain, 65)

Anterior Cingulate Gyrus and Learning

Anterior cingulate gyrus is well connected with the amygdala and other structures of the limbic system that regulate our emotions, the fight-or-flight mechanism, and conditioned emotional learning. (Ratey; User's Guide to Brain, 165)

Feedback Maximizes Learning

Studies suggest that feedback is necessary to maximize learning.  The brain is exquisitely designed to operate on feedback, both internal and external. (Ratey; User's Guide to Brain, 179)

Substantia nigra is critically involved in the feedback process of learning. (Ratey; User's Guide to Brain, 179)

During the feedback process, what is received at any one brain level depends on what else is happening at that level and what is sent to the next level depends on what is already happening at that level.  The brain is self-referencing, which allows interactions to provide constant feedback, crucial to our ability to learn. (Ratey; User's Guide to Brain, 179)

Memory and Learning

Memory is a central functionality that brings together learning, understanding, and consciousness. (Ratey; User's Guide to Brain, 185)

A memory is only made when it is called upon.  In its quiescent state, memory is not detectable.  We cannot separate the act of retrieving and the memory itself. (Ratey; User's Guide to Brain, 186)

Memory is not fixed at the time of learning but takes considerable time to develop its permanent form.  Fixation process. (Squire & Kandel; Memory, 102)

Learning is of a Hebbian nature such that simultaneous pre- and post-connection activity leads to increased connection strength    and is often used in explicit analogy to synaptic processes such as long term potentiation (LTP). Andersen; Hippocampus Book, 716)

Relational processing at encoding enables flexible access to information and situations    quite different from those of the original learning. (Andersen; Hippocampus Book, 662)


Research study — Retrieval of Knowledge Enhances LearningNot only does retrieval produce learning, but a retrieval event may actually represent a more powerful learning activity than an encoding event.


Bits and pieces of a single memory are stored in different networks of neurons all around the brain. (Ratey; User's Guide to Brain, 186)

Formation and recall of each memory are influenced by mood, surroundings, and gestalt at the time the memory is formed or retrieved. (Ratey; User's Guide to Brain, 186)

Memory changes as we change over time. (Ratey; User's Guide to Brain, 186)

Each memory arises from a vast network of interconnecting pieces. (Ratey; User's Guide to Brain, 186)

A memory is a tiny bit different each time we remember it. (Ratey; User's Guide to Brain, 186)

Frontal cortex is the part of the brain that organizes the bits and pieces of a memory into a temporal, logical, and "meaningful" story. However, it must be set in motion by the amygdala, which provides an emotional tag to a memory, a "meaning" that helps cement the pieces. (Ratey; User's Guide to Brain, 186)

Antonio Damasio’s "Convergence Zones"

Antonio Damasio proposes "convergence zones" that oversee the collection of the names of objects and animals and other zones that unite sensory information about people, perception, and emotion. (Ratey; User's Guide to Brain, 187)

Convergence zones enable us to automatically conceive of objects, ideas, or interactions as a whole, if the pieces have been put together enough times. (Ratey; User's Guide to Brain, 187)

Damasio also proposes that there is a hierarchy of convergence zones.  Lower convergence zones link to cues that allow us to understand the general concept of a "face" while higher convergence zones allow us to recognize specific faces.  Linking the two are intermediate convergence zones that differentiate details in individual faces -- nose line, pallor, eye shape. (Ratey; User's Guide to Brain, 187)

Hippocampus and Memories

Hippocampus does not store memories.  It has been likened to an intelligent collating machine, which filters new associations, decides what is important and what to ignore or compress, sorts the results, and sends various packets of information to other parts of the brain.  It is a way station that hands out the pieces. (Ratey; User's Guide to Brain, 188)


Research Study — Hippocampal Place Cells and Long-Term Memories


Sleep Is Important for Memory

Sleep, associated with dreaming, is important to human memory. (Ratey; User's Guide to Brain, 188)

Brain Forms Concepts

When the brain forms concepts, it constructs neuronal maps of its own activities.  The neuronal maps categorize, discriminate, and recombine the various brain activities needed to form ideas and emotion. (Ratey; User's Guide to Brain, 189)

We need and use certain types of memory more than others; our brains can extend the regions response for specific functions, even recruit new regions to help. By exercising our brains we can strengthen our memory. (Ratey; User's Guide to Brain, 190)



Return to — Memory

Link to — Consciousness Subject Outline

Further discussion — Covington Theory of Consciousness