The Music Lesson by Lord Frederic Leighton (19C British pre-Raphaelite)

Learning as a biological phenomenon
"...observations of natural learning tend to encourage the view that learning consists, not of a unitary general capacity, but of a collection of specialized abilities which have evolved independently in particular species in order to do specific jobs."

Reflexes and "instinctive" behavior are good ways to respond rapidly and reliably -- especially in a reliable environment. But environments and organisms are always changing, so BEHAVIORAL PLASTICITY can be good thing. Learning is an expression of plasticity. Basic behavioral patterns are elaborated upon or combined in varying ways to better cope with the changes in and around us.

As the proximate cause of behavior, and the key system changed as result of experience, we should have a perspective on neuroplasticity.

Learning is among the most plastic of all phenomena, allowing organisms to adapt to the unique properties of the environments in which they find themselves. The competence to learn specific associations varies within limits (constraints) that have themselves evolved.

Learning as a biological phenomenon

"...observations of natural learning tend to encourage the view that learning consists, not of a unitary general capacity, but of a collection of specialized abilities which have evolved independently in particular species in order to do specific jobs."

When asked "what is learning for?" T.J. Roper. believes it that involves information pertaining to the individual's experience in the world rather than experiences so reliably encountered by the species that these needs would have come to be encoded in the genome. (1983 in Genes, Development, and Learning, Halliday and Slater, eds. pp 178-212.) feels it is to solve practical problems. Galef (1976 in Adv Stud. Behav, 6:77-100) One of the practical problems is particular to social species: the need to make predictions about conspecifics (A. Jolly, 1966, in: Science 153:501-506; N.K. Humphrey, The social function of intellect, in Growing Points in Ethology, Bateson & Hinde, eds, pp 303-317.)

Arguably, the need to make predictions about other individuals is one of the arguments for the evolution of consciousness (first mentioned (as far as I know) by Thomas Hobbes).

Learning Defined. We may define Learning as a "change in behavior as a result of experience" but in practice how are other causes of changes in behavior --such as maturational changes-- identified?

In non-associative learning an animal changes its "response to a stimuli without association with a positive or negative reinforcement.

-- Habituation (diminished responsiveness). Animals frequently subjected to a stimulus will often become habituated to that stimulus--they will show a reduction or total elimination of response to a stimulus without positive or negative reinforcement. If you poke them, sea slugs (Aplysia) will curl inwards. However, if you poke them repeatedly, the response will become less and less extreme until they do not withdraw at all. When presented with a novel stimulus, such as an electric shock, the sea slugs will recover their withdrawal response to poking. This phenomenon in which the habituation disappears is, conveniently, known as dishabituation.

--Sensitization (enhanced responsiveness) "...Sea slugs can [also] be sensitized, whereby they will show an increased response to poking after first being presented with a strong or novel stimulus. The difference between dishabituation and sensitization is that dishabituation involves the recovery of the original response while sensitization produces a response stronger than the original one.

  • non-associative learning is often associated with drug-dependency (more)

In associative learning an animal learns the relationship between stimuli and behavior

-- Classical Conditioning (the animal makes an association between two stimuli) (in 1902, the Russian physiologist, Ivan Pavlov, began his famous experiments on conditioning. Pavlov repeatedly presented a dog with food following the ringing of a bell. When the bell sounded without the presentation of food, the dog would still respond to the bell as if it were food. Pavlov collected the dogs' saliva and found that the amount of saliva produced by bell ringing increased as the dogs were more frequently exposed to the coupling of food presentation and bell ringing. The dog had learned to associate the sound of the bell with food. Pavlov called the food an unconditional stimulus, or UCS, because the dog's normal reaction would be to salivate at the presentation of food. The bell he termed the conditional stimulus, or CS, because response to the bell was conditional upon the association between the bell and food. For the same reasons, salivation in response to food was labeled the unconditional response, or UCR, while salivation in response to the bell was called the conditional response, or CR. Conditioning the dog to salivate at the sound of the bell occurred as a result of a contingency between the UCS and the CS. Pavlov's experiment was an example of positive conditioning. It is also possible to negatively condition an animal by using an unpleasant UCS.)

-- Operant Conditioning -- (the animal learns about the relationship between a stimulus and the consequences of its actions) (In classical conditioning, receives no benefit from associating the CS with the UCS. However, in operant conditioning, an unassociated behavior becomes associated with a reward. B.F. Skinner designed an apparatus called a "Skinner box" to test the interaction between UCS and CS. A rat was placed inside the Skinner box; if the rat pressed down a lever inside the box then the box would release a food pellet. Soon, the rat pressed the lever far more often than he would just by chance. Most likely, the first time the rat pressed the lever it was by chance. But with each instance of lever pressing, the operant is reinforced by reward with food. The rat learns that pressing the lever is associated with food, and so he will increasingly press it. Almost any operant and reward system can be used effectively.)


Human experience is based on learning that our actions affect subsequent positive or negative outcomes. Rewards strengthen associations between contextual stimuli and actions thereby reinforcing and maintaining successful behavior; whereas punishments induce avoidance of erroneous actions. While we usually learn from both positive and negative reinforcement, the relative amount of learning from success or errors varies between individuals. Klein et al. (2007) investigated a human genetic polymorphism associated with the density of brain dopamine D2 receptor. Reduced D2 receptor density was associated with less efficient learning from errors. In people with lower D2 receptor density, the reduced capacity to learn from errors was accompanied by reduced feedback-related activity in the posterior medial frontal cortex, an area known to monitor for negative action outcomes.
(Science vol 318:1521 "Learn from your Mistakes") (more)
-- Insight Learning -- involves the combining of isolated experiences into a new experience that is effective for gaining a desired result. The new experience often occurs suddenly and spontaneously as in an "Aha!" experience. It appears to happen in some animals but is hard to identify with confidence.
    (Wolfgang Kohler was a Gestalt psychologist who used chimpanzees in the study of insight learning. His most famous observation was of a chimpanzee named Sultan, who had learned to use a stick to reach bananas just out of reach. When Kohler placed a banana too far away to reach with just one stick, Sultan used two sticks. After a period of playing with the sticks, he suddenly fitted the two sticks together and was able to reach and recover the banana. Kohler used the term "insight" to describe Sultan's apparent sudden “perception of relationships.” Another interpretation is based on the fact that chimps play a lot: Michael Chance observed chimps playing with boxes and one time piling them up so they could almost reach the top of their cage. Combined with a reward, it seemed trial-an-error learning was combined with "learning sets" to lead to insight. )

Learning Sets -- involve organized patterns of responses appropriate in one context that can be applied to a new, more complex problems.
IMPRINTING ... irreversible and limited to a critical" or "sensitive" period in an animal's life ...; related to MATURATION of INPUT, INTEGRATIVE, or OUTPUT COMPETENCE? (output can be important in particular because the feedback to the integrative functions that would cause the behavior must be reinforced to continue developing. (what happens if a song-bird is deafened at an early age?)
  • Fish imprinting? some cichlid fry will approach another fish indiscriminately when they are up to 4 days old, but between 4-16 days they are selective for their own species- seems a balance between imprinting and innate species preference
  • Ducklings- sensitive period for imprinting is sharply between 13-16 hours posthatch, declines to zero by 35 hours. in open-nesting species, vocalizations facilitate the response; also, isolates respond better than ducklings placed in groups
  • Ring Doves, reared from eggs by graduate students at Rutgers revealed only after sexual maturation that they had "sexually imprinted" on the hands of the care-givers.

"PREPAREDNESS" resembles "readiness to learn" (which involves a combination of growth,such as maturation of nervous system or sensory organs and experience,such as reading requires a previous background of spoken words and experience with letters and pictures).

Preparedness, however, emphasizes the genetically programmed disposition to learn some things more easily or with less experience than others. Biological make-up "prepares" organisms to make certain associations, while they are "contraprepared" to make others. This has been applied to various phenomena such as the apparent evolutionary basis of specific fears and to "conditioned taste aversion" ("bait shyness") in which organisms learn to avoid specific foods after getting violently sick after eating/drinking them: it can take only one trial, the experience and its consequences can be separated by hours.

The phenomenon was researched by John Garcia and developed further as "biological readiness to learn certain associations because of survival advantages" by Martin Seligman.
  • SOCIAL learning - Emphasis is on OBSERVATION and MIMICKING ... latest findings on MIRROR NEURONS underscore the evolved mechanisms (which when dysfunctional may lay behind several important behavioral dysfunctions, such as sociopathy and autism).

New models of the manner in which the stress response affects behavior includes the influence of subclinical stress which can be evoked by any mismatch between internal models of reality and the apparent external world (see cognitive dissonance).

Jean Piaget, one of the founders of the modern understanding of child development regarded assimilation and accommodation as the central processes of the development of knowledge.

the resolution of COGNITIVE DISSONANCE is seen to be a key "energizer" of learning and "reconfiguring" information already possessed [more on cognitive dissonance, including classic mechanisms]

When there is a mismatch between ACTIONS and INTENTIONS (the animal does not get a desired result) cognitive dissonance occurs (perhaps in proportion to the importance of the act), and the organism is in some measure stressed. Internal models are based on countless experiences in which new information is assimilated into the model (set of expectations) or accommodated (the model changes slightly). In the stream of ongoing experience, the world is always being "tested" to see that expectations are more-or-less likely to be met ("reality testing") stimuli that meet certain criteria are also "tested" to see if they "fit in" with preceding experiences that they more-or-less resemble ("establishing coherence"). When stimuli do NOT meet the tests (which are highly automatized and always ongoing) the organism's attention is refocused on the "anomaly" -- the unexpected experience. The internal model is challenged and the level of attention is enhanced, possibly even up to conscious investigation.
HUMAN learning cycle and "alternative styles" (Kolb)


LATENT LEARNING [more on latent learning]

CONSTRAINTS on LEARNING -- [more on constraints]

MEMORY involves an organism's retention of information, usually based on individual experience. Affects on their quality are primarily Encoding (processing and combining of received information); Storage (creation of a permanent record of the encoded information), and Retrieval/Recall (calling back the stored information in response to some cue for use in some process or activity) [more] [memory in Wikipedia]

    Representation of past experience of which we may or may not be conscious; each unique memory is likely a unique pattern of neural activity, reactivated when called for, but potentially vulnerable to neuroplastic changes. The several forms of memory are each mediated by different patterns of wiring often associated with different regions of the brain. Two forms are often distinguished, short term ("working") memory and long term memory (2 forms: PROCEDURAL and NON-DECLARATIVE) that remain accessible after years. [more]



We NEED two apparently dissimilar things in life: the MAINTENANCE OF STABILITY, the status quo,a circumstance of past success and safety, and GROWTH -- an enlargement of knowledge and competence, of biological potential. Stability and (more conspicuously growth) are epigenetic processes in which genes and experience interact to help the organism meet present and future needs. Much growth is circumstantial, unintended, and much is intentionally acquired. Reasonably, the ratio of stability to growth changes as we mature and meet different needs. The "tension" between these functions was characterized as the "essential tension" by historian/philosopher of science, Thomas Kuhn: he represented them as "tradition" and "innovation." Cognitive Dissonance is a conscious or nonconscious perception of a mismatch between the stable or active circumstances we find ourselves in and those in which we intended to experience. If a corrective to circumstances fails or is likely to fail, we often rationalize or reconfigure our interpretation to relieve the tension generated.

HUMAN DEVELOPMENT -- "The Scientist in the Crib" "To most of us, tiny babies don't appear to know very much. But not to Alison Gopnik, professor of cognitive psychology at the University of California at Berkeley. After embarking on a career in philosophy of science, she switched to studying babies just as developmental psychology was taking off. She has written about some of the unexpected discoveries in the recent book The Scientist in the Crib, which explains the huge changes in our view of babies over the past thirty years" [more]

PHYSIOLOGY of LEARNING -- CNS and endocrine systems are the PROXIMATE CAUSE of behavior and can be studied at the levels of

    OPTIONAL: LOOKING more deeply: "Natural rewards preferentially stimulate dopamine transmission in the nucleus accumbens shell. This effect undergoes adaptive changes (one-trial habituation, inhibition by appetitive stimuli) that are consistent with a role of nucleus accumbens shell dopamine in associative reward-related learning. . . . Addictive drugs share with natural rewards the property of stimulating dopamine transmission preferentially in the nucleus accumbens shell. This response, however, in contrast to that to natural rewards, is not subjected to one-trial habituation. Resistance to habituation allows drugs to activate dopamine transmission in the shell non-decrementally upon repeated self-administration. It is hypothesized that this process abnormally strengthens stimulus-drug associations thus resulting in the attribution of excessive motivational value to discrete stimuli or contexts predictive of drug availability. Addiction is therefore the expression of the excessive control over behaviour acquired by drug-related stimuli as a result of abnormal associative learning following repeated stimulation of dopamine transmission in the nucleus accumbens shell."
Di Chiara G. 1999. Drug addiction as dopamine-dependent associative learning disorder. Eur J Pharmacol. 375(1-3):13-30.

CONFIDENCE in BELIEFS, VALIDITY, and "TRUTH" -- Among the premier (sole?) adaptive function of learning is the attaining of valid information about the world upon which our intelligence may then act (for example to develop and "experiment" with a mental model to better understand the consequences of potential actions) --Such valid ideas are regarded as "true" -- but truth has at least two key dimensions: Correspondence and coherence.
"People's beliefs have some measure of validity (external correspondences) and reliability (internal coherence). A more intelligent, adaptive person has achieved a higher degree of external correspondence and internal coherence in his or her knowledge based and belief structures. People think unintelligently to the extent to which they make errors in achieving external correspondence or internal coherence. (Sternberg, Robert J. (1997) "The Concept of intelligence and its role in lifelong learning and success". Amer Psychol., 52(10):1030-1037).

    IT is important to have CONFIDENCE in the VALIDITY of what we learn -- DO OUR senses deceive us? Will specific stimuli reliably indicate specific experiences? will specific actions we might take in response to a stimulus have a reliable outcome? What is the "truth?"
    The principal dimensions of "truth" are
        • Percepts "correspond" to their sources in the "real world"
        • relies heavily on our curiosity
        • it is basically inductive ("particulars to generalities") and
        • it provides the premises of our life-theories (on which logic may then act)
        • as well as the anomalies that challenge them.
      and COHERENCE
        • Percepts "cohere" ("fit in with") other percepts
        • relies heavily on our story-telling ability
        • it is basically deductive ("generalities to particulars") and
        • strengthens our confidence in life-theories by
        • reconciling internal contradictions and accommodating anomalies.
    confidence and effectiveness of action are correlated - but confidence wanes when anomalous correspondences are detected
When is enough? : As an evolving organism, is there a point at which one could say, "my fitness is maximized, I need reach no further."

MORE RECENT INSIGHTS ABOUT HOW WE LEARN from early childhood development experts: Alison Gopnik did a nice job making a favorite point of mine: "that schools don't teach the same way children learn." She cites Barbara Rogoff for observing "guided discovery" in Guatemalan Indian villages: "youngsters gradually mastered complex skills . . . by watching adults, trying themselves and receiving detailed corrective feedback about their efforts." Rogoff contrasted guided discovery and routinized learning. This got my attention because it echoes what I propose about the neurobiology of discovery versus invention (correspondence versus coherence). Gopnik wrote that "the two modes of learning seem to involve different underlying mechanisms and even different brain regions, and the ability to do them develops at different stages. Babies are as good at discovery as the smartest adult -- or better. But routinized learning evolves later." Read "How we Learn"

CAN WE learn anything from the "artful scribbles" of children as their representations change? visit the "YOUNG at ART website [link lost]

DEVELOPMENT: HOW MUCH of change with time is attributable to ONTOGENESIS --"simple" maturation of input, integration, and/or output competencies -- and how much is learning. Capacity to learn changes as the organism's brain changes and as the experiences change. Often there are "sensitive periods" for learning specific kinds of associations. Changes in humans are among the most documented: one emerging point of view is that of Alison Gopnik's "scientists in the crib" [more]

Relationship between LEARNING and TEACHING -- there is a vast reservoir of research and information about the biology of learning but very little on the biology of teaching:
A systems model of teaching-learning relationships involves INPUT, INTEGRATION, and OUTPUT

from Greenberg, 1991
    TO LEARN, information from the body (SOMATIC) or the ENVIRONMENT (including information from the TEACHER) must be DETECTED by the sense organs (INPUT), must be integrated into the pre-existing structures of the CNS (INTEGRATION), and it may then result in a detectable CHANGE IN BEHAVIOR (OUTPUT). [more on teaching/learning]

    It is possible to learn without consciousness or memory of the learning experience -- in "latent learning" the OUTPUT (change in behavior) is not manifest until a specific eliciting stimulus is experienced. Such latent knowledge can affect the expression of other behavioral patterns (in some views, psychoanalysis is a search for latent or subconscious knowledge that affects the expression of other behavioral patterns in dysfunctional ways; other views see art as an expression of subconscious knowledge, sometimes in very compelling ways and sometimes in ways that can provide useful insights for other people) [more on latent learning]

LOOKING AHEAD TO PHYSIOLOGY as a approach to insight about development

The structure and function of the nervous system is generally presumed to be the materialist basis for all behavioral actions. examples of neuroplasticity

LOOKING AHEAD TO ECOLOGY as a approach to insight about development
Cognitive Ecology

Internal and environmental information converge to shape behavior, but the capacity to change as a result of experience and the paths change may take are highly subject to environmental selection pressures, guiding evolution of traits that can best be characterized as cognitive. “Cognitive Ecology lays the foundations for a field of study that integrates theory and data from evolutionary ecology and cognitive science to investigate how animal interactions with natural habitats shape cognitive systems.” Essay on evolutionary ecology of learning (master idea of "phenotypic plasticity")

EC; What are the first few opening lines of Aristotle's METAPHYSICS. Relate those lines to the idea of BIOLOGICAL NEEDS and the view that perception is intrinsically pleasurable as described in "Perceptual Pleasure and the Brain"

LEARNING MAY or MAY NOT involve a teacher!
sometimes the LEARNER is the TEACHER [more]


Education and the brain sciences from the "21st Century Learning Initiative" --


Oct 2010