Evolutionary Cognitive Neuroscience MacNeilage Assignments

PSY 341K - Evolutionary Cognitive Neuroscience

MacNeilage: Evolution of Speech. Chapter 1. Summary

Best explanation of speech is in terms of the “ultimate causes” of evolutionary biology (Mayr).

Dobzhansky: “Nothing in biology makes sense except in the light of natural selection”
(Though note culture).

Darwin: Survival criterion: successful use (Action) For speech, the survival criterion is successful social communication.

Thus mind (cognition) arises from body use. The monograph topic is Movement-Mind relationships, following Sperry: Infer mind from movement. The rationale

: 1. Movements are readily available for observation. Inferences from movement to mind are relatively simple, compared, for example to grammar.

Note Lashley and the serial order problem.

But action hasn’t historically been part of mind. Several reasons.

The Neodarwinian Perspective (p 7)

Roughly it is “Survival of the Fittest” (Spencer)

All life derives from a single form. p 8. The “Secret of Life” (Crick)

Note here the role of self organization, but always plus selection.

The problem of “Just so stories”.

But some evolution is observable: Darwin’s and the Grant’s finches. Galapagos Island finches beak shapes change over time with food availability.

Note the tinkering metaphor.

Note also “exaptation”: Example: Evolution of the speech apparatus.

Inevitable messiness results from tinkering.

Darwin’s basic tenet: “Descent with Modification”.

Jacob: Two ways: Transformation and combination.

What has been hard for people to stomach is that the Neodarwinian perspective applies to humans as well as other animals. Two main reasons:

1. We are definitely very different.
2. Religious and philosophical beliefs that we are a breed apart.

Cartmill et al.: Emphasizing human specialness makes solving the problem of descent with modification harder. Note the continuity paradox. Nevertheless we must bite the bullet and come up with a modification scenario for humans.

The Classical Perspective

Russell: Basic philosophical theme: The search for permanence in change.

One ‘solution’ immortal God (Woody Allen’s preferred solution).

600 B.C.: Heraclitus emphasized the importance of change: You can’t go into the same river twice.

But by 400 B.C. the notion of permanence ruled and it has ruled ever since. This was a bad development for the discipline of history. General principles sought.

The downgrading of history was not surprising given how little of it was available to the Greeks. But a second development supporting permanence was the rise of mathematics.
Pythagoras’s work was the first key development.

Rationalism: Thought and reason were considered the main source of knowledge. E.g. Geometry: Axioms plus deduction.

Main Greek Philosopher: Plato. Believed in underlying forms (“essences” hence “essentialism”). E.g. triangle. Note Russell on beds.

Formalism: Concern with underlying forms.

Main Renaissance philosopher: Descartes: Applied Cartesian doubt. Geometricians as models p 17 End point: cogito ergo sum (I think, therefore I am)

Two key dualisms: 1. Mind/Body; 2. Humans/Animals We share the body with other animals, but only we have mind.

Ideas considered innate. Quote, p 18

Triangle considered a key concept, though note Hebb’s observations.
Essentialism and Neodarwinism are incompatible. A priori forms cant be modified. Note Dennett quote, p 19

Relevance of the classical perspective to speech: Noam Chomsky’s “Cartesian Linguistics”. He notes that for Descartes, language is criterial for mind. Quote, p 20.
For Chomsky, the creativity of language is crucial: It has unbounded scope and is stimulus-free.

Chomsky’s focus: Generative grammar. Quote, p 21. Note the competence-performance distinction. Competence is the linguistic component of the Cartesian mind.

Basic concept: Universal Grammar (UG). It has two components: 1. Syntactic; 2. Phonological. Chomsky adds phonology to Descartes’ mind. He describes language acquisition as solving Plato’s Problem.

Comparing the two approaches.

Darwinism: Mind (cognition) derived by descent with modification based on successful action across the time domain (generations). Variation is essential as the raw material for selection.

Classicism: Human mind has one eternal fixed form for all. Thus both the time domain and variability are irrelevant.
Plato and Descartes: Origins problem solved by Gods. Chomsky: Origins problem solved by “a mutation” (but he doesn’t believe in natural selection). Quote pp 24-25. He’s a closet essentialist.

One problem to be solved: Mathematics must pay the cost of its irrelevance to nature.
Its success comes from its allure. See Russell quotes, p 25. Crick: Physics and biology are different p 26. Plotkin: Psychology has suffered from its physicalistic (mathematical) “now and forever” bias.

Problems of explaining speech from the classical viewpoint.

Tinbergen: Four questions to be answered in order to understand any biological phenomenon:
1. How does it work? (Function)
2. What does it do for the organism (Purpose)
3. How did it get that way in phylogeny (Evolution)
4. How did it get that way in ontogeny (Development)

The time domain is essential to all of these questions.

Q 1.
Here is where Lashley’s Serial Order Problem comes in.
His conclusion for language: “The order must be imposed upon the motor elements by some organization other than direct connections between them.”
This was a Chomsky-like alternative to the behavioristic notion that serially ordered behavior is chains of reflexes. p 29. Millwright question.

For Lashley the order of language came from a level between the prelinguistic thought that lies behind a piece of language output and the movements made to speak it.

Unlike Chomsky, he was trying to get from mind to movement.

Lashley made two other important contributions:
He anticipated the concept of working memory.
He emphasized the importance of rhythm generators in achieving serially organized output. Note the importance of the syllable.
None of this is of any importance to Chomsky because it is in the performance domain.
But, as Thelen and Smith assert: “If competence in the Chomskyan sense is part of our biology, then it must be embodied in real-time processes.” Thelen and Smith, p 27.

Q 2.
Speech enables us to send and receive a potentially infinite set of messages.
Jakobson: We speak to be heard in order to be understood.”
Crucial concept here: Minimal pair. It allows distinction between words.
More generally, efficient message transmission is thought to be achieved by a continuously present trading relation between ease of production and perceptual distinctiveness. This trading relation is necessarily worked out in the time domain:
This is all irrelevant to Chomsky who believes that the contribution of language when it first appeared was to facilitate thought.

Q 4.
Time domain taken for granted by Chomsky. The main approach is to note phenomena that don’t appear to be learnable and then to assert that they must result from a genetically specified UG; e.g. error correction. Note “Plato’s problem: the poverty of the stimulus. Note Tomasello’s “cut and paste” alternative.

Also there is much agreement that genes underspecify behavior.

Note Stent: “too many removes’.

Dawkins: “blueprint” and recipe metaphors.

Chomsky: “the argument from ignorance.” Note Helmont’s error.
Chomsky is missing culture. Donald: “Language does not self-install.”
Note the “E-language”-“I-language” distinction.

Culture: “Shared knowledge and beliefs.” Plotkin. “Memes.”
Genes are in the environment. Cosmides and Tooby quote, p 38.
Chomsky’s conclusion of simple genetic determinism inappropriate.
Note Plotkin quote, p 39. And note Fig 1.2.

Circularity and the mist-clearing metaphor for development. What determines the landscape. “Markedness” is circular. Miller: Linguists tend to accept simplifications as explanations.”

Specific problem here: Kids are not just little adults. The priorities of early speech are superceded.

Q 3.
Considered earlier. Ahistorical approach to language unrealistic.

An Emerging Intellectual Context: “Embodiment.”

Note Lakoff, Johnson, Clark, Damasio, Edelman, Dynamical systems perspective-e.g. Thelen, (with self-organization. Note hexagon example). Wilson (Robertson-Davies: “The hand speaks to the mind as surely as the mind speaks to the hands.”)

“Evolutionary Psychology”. I don’t like the emphasis on innate modules. It tends to finesse Tinbergen’s questions 3 and 4.

Plan of the Book.

Notes from P. Tort: Darwin and the Science of Evolution.

Chapter 1. An Uncertain Beginning

Family background: Erasmus Darwin: “Zoonomia” touted evolution.
Also Wedgewoods.
Undistinguished university career, but collected a lot of beetles.
Influenced by Paley: Natural Theology.

Chapter 2. The Voyage of the Beagle

Henslow plus Wedgewood pointed him towards the Beagle (Left on Dec 27, 1831 with the first volume of Lyell’s Principles of Geology.
Cape Verde. Observed Lyell’s uplift-subsidence cycle.
Lyell’s volume 2 introduced Lamark. He believed in spontaneous generation of life and in evolution (though of acquired characteristics. Used modifications under domestication as evidence.
Politically, he got the worst of Cuvier and “catastrophism”.
Lyell: first modern use of “evolution”. Didn’t like Lamark because he himself believed in “uniformitarianism”. Note also “actualism” and “present causes”. Changes under domestication is an actualist finding.
Interacts with savages in Terra del Fuego.
Various evidences of earth upheavals in South America: not evidence for catastrophism. Instead found evidence for gradual changes, including gradual extinction.
Much interesting stuff here on biogeography.
Geological evidence suggested the omnipotence of time.
15th of Sept. Galapagos.
Noted 13 finch ‘varieties’, later to be designated as species.
They were native to the islands but related to mainland finches.
Formulated his coral reef theory.
Arrived home 2 Oct 1836
Specimens entrusted to various specialists.
Sept. 28th, 1838: He realized the implications of Malthus’s theory. Population outpaces food. Something’s got to give. Natural selection determines what wont give.

Chapter 3. The Theory of Natural Selection.

He focused on establishing variation. Sent out a questionaire to stock breeders. Concluded that nature behaves like breeders do.
Got married and eventually withdrew from public life to Down House.
Chronically bothered by Chagas’s Disease.
Wrote two unpublished monographs outlining his theory in 1842 and 1844.
Published Geology of the Voyage of H.M.S Beagle in 1846.
Worked for 4 years on a monograph on cirripedes, including barnacles. Establishing reputation.
Interested in classification difficulties as problems for creationist and earlier typological conceptions.
Also interested in metamorphoses as evidence that change rather than ‘fixity’ is the law of the organic world.
Had 10 children. Three died. Death of one finally disillusioned him regarding religion.
Alfred Russel Wallace publishes an article entitled ‘On the law which has regulated the introduction of new species.’
At Lyell’s instigation he began working on what became Origin of Species.
June 1858: Darwin receives Wallace’s manuscript: On the tendency for varieties to depart indefinitely from the original type.
Friends rapidly organized a joint presentation of Wallace’s and Darwin’s stuff to the Linnean Society on 1 July 1858.
Darwin, who had scads more evidence and arguments prevailed.
Wallace did not go ‘the whole orang’ regarding humans as different and embracing spiritualism.

Chapter 4. A Resounding Triumph.

Origin of Species published on 24 Nov. l859. 1250 copies sold out that day. Quote on content, p 76.
Only 1 sentence on humans: “Psychology will be based on a new foundation, that of the necessary acquirement of each mental power and capacity by gradation. Light will be thrown on the origin of man and his history.”
1871. Anthropological questions addressed in The Descent of Man.
In the interim, Spencer and Galton confused the issue.
Spencer: Social Darwinism: “social philosophy that was ultra-liberal, individualist and anti-state, strongly in favor of competititive conditions, and fiercely hostile to the concept of aid to the poor.
Galton: Made some contributions to mental measurement but advocated eugenics.
1860: Big confrontation at the Oxford meeting of the British Association for the Advancement of Science. Wilberforce put down.
In the 1860s Darwin worked on The Variation of Animals and Plants under domestication.
He added a hypothesis about “gemmules” for hereditary transmission.
But stuck with the problem of blending inheritance. Fleeming Jenkin.
Other criticisms. Why no intermediate forms? Incomplete fossil record, and new forms replace older ones.
Kelvin: Earth too young judging by its heat. Underestimated radioactive effects.

Chapter 5. Nature and Civilization

Quote p 93 on highest part of man’s nature.
The biological advantage has become a social advantage.
Emphasized the moral superiority of women.
1872: The Expression of Emotions in Man and Animals
3 principles: Useful habits, antithesis and direct action of the nervous system. Argument for evolutionary continuity.
Planned ‘Big Book on Species’.
1876 Autobiography
1881 book on worms.
Died on April 19, 1882.
Buried in Westminster Abbey with Henry V111, Newton & (now) Churchill.
On religion: “disbelief crept over me at a very slow rate.
“He contrasted the infinite suffering of living beings with the image of an omnipotent and benevolent God.
“put the value of religion into perspective by showing that the peoples of the world all have different beliefs and traditions.
But never declared that God did not exist.
p 107: A living theory of civilization.

Notes on Stanley, S. The New Evolutionary Timetable.

Chapter 2. The Voyage Towards Evolution

Lyell’s uniformitarian message to Darwin. The earth was extremely old.
(Hutton: “no vestige of a beginning, no prospect of an end” p 190
If geology is changing why isn’t biology changing in concert. If biblical origins story isn’t true, why believe it on the matter of life’s origins.
Earthquakes not catastrophism but Lyellian gradualism.
Cuvier: Correlation of the parts implied for him perfection of design, and therefore immutability (except for catastrophies). Note though that there were mass extinctions.
His thinking helped the evolution idea in two ways: 1.his “progressionism” – each new wave is better – helped pave the way for evolution. 2. His 4 “embranchments”, in contrast to Aristotle’s great chain of being (note also plenitude) encourged people to think of branches.
Lyell believed plenitude. While recognizing extinctions, he thought the consequence was “expansion of other species, or .. the appearance of altogether new but similar ones.” (p 23. He didn’t see progression in the fossil record and encouraged Darwin to believe in its gaps.
The voyage made Darwin believe in evolution. Natural selection came later.
What took him from creationism to evolution. Much of it was a set of geographical arguments. One thing was the apparent irrationality of creation. Same geographical conditions in different parts of the world but somewhat different biotas. There were not, therefore multiple origins of the same thing in different places” “single place of origin” rule.
He noticed also that widely dispersed species had the power of dispersal.
Despite the above differences, similarities (e.g. ostriches, emus, rheas) suggested single origins. Why a single body plan for related nearby species, but with changes in habits (ground & tree sloths).
Why such differences on two nearby sides of geographical barriers (e.g. Atlantic and Pacific sides of Panama.
Why such impoverished biotas on islands. Only mammals on some islands are bats who could have flown there.
Why so many species endemic to the Galapagos, especially land birds.
Why one turtle variety per island.
Conclusion: “Species endemic to newly formed islands were derived from nearby land masses by some natural agency. Only in this manner was it possible to explain why the young insular faunas resembled nearby continental faunas in general but were unique in detail.” p 34. Darwin: “such facts would undermine the stability of species.”

Chapter 3 The Origin and its Very Slow Process

With Darwin, classification went from chain to tree with its attendant hierarchical connotation.

Embryological evidence also encouraged the evolutionary idea. Common early form suggests common origin.
Another interpretation: An animal passes through the scale of being during development. But contra Haeckel, no adult forms.
For Darwin, why would a creator begin with a simple common mold for species that are later so distinct? For him if very different breeds of dog started with a common embryological pattern, why didn’t natural selection work this way too?
Why was the creator so wasteful as to produce all those embryological vestiges?
Why homology – same bones for wing and leg of a bat?.
Owen handled some of this with the concept of God-given “archetypes.
Problem was the fossil record didn’t give the gradualism scenario, so Darwin emphasized its problems.
Natural selection confronted the Platonic ideal.
Had a problem in convincing people there was sufficient variability. Chapter 1 title is indicative of his efforts “Variation under domestication.”
Problem: variation denied a priori by Essentialism.
September 28, 1938, (on my birthday) he realized the significance of Malthus’s arguments for his thesis. p 43. He transferred species struggle to individual struggle.
Secondary devlopment: sexual selection. Male characteristics are hardly consistent with the creationist notion of perfection of creation.
“Evolved” (evolution) appears only as the last word of Origins.
(Note that the phrase “Natura non facit, saltum” – “Nature does not make jumps” came from Scala Naturae times, but was embraced by Darwin.
He emphasized gradualism partly because he worried that people would not believe his thesis. Also he tended to polarize gradualism as the opposite of instantaneous creation.
Also gradualism tended to get him off the hook with respect to any assumption of perfection. He had to deny perfection but argue that selection could produce things like the eye that had claims to being perfect.
Darwin didn’t say much about the speciation process and didn’t capitalize on allopatric speciation a la Mayr.
He was bedeviled by the notion of blending inheritance which would tend to move offspring towards the mean of their parents.
Problem: life is so tightly packed according to the plenitude idea that little variability must be tolerated. Another motivation for his gradualism.

Chapter 4: Darwinism Challenged, Darwinism Affirmed.

Evolution has fared well but selection has had its ups and downs.
Darwin’s strategy was to claim opponents as partial supporters thus making it more difficult for them to maintain their opposition.
He also made efforts to win over Lyell, Hooker and Huxley.
Lyell eventually tilted from immutability to evolution.
Hooker was an “unwilling convert” to natural selection.
Huxley became convinced of evolution but not natural selection.
The main support of these 3 was for Darwin’s good science.
Two reasons for Darwin’s delay of publication:
1. He felt he had to establish his reputation first. So he worked on barnacles.
2. He had to demonstrate within-species variation. He corresponded with breeders and worked on pigeons.
Note: 1868 “Variations of animals and plants under domestication.”
Also he was deterred by the religious opposition.
Problems with acceptance:
Kelvin: the world was too hot to be older than 20mya. He underestimated radioactive effects.
Heredity: Blending inheritance produces regression to the norm. (Fleeming Jenkin).
Mendel’s book on particulate inheritance remained uncut in Darwin’s office.
End of 19th century saw a movement to experimental biology in search of the prestige of the physical sciences.
But Mendel’s contribution led to an overestimation of the contribution of mutations (note particularly macromutations) and a belief that selection was a more minor effect. Speedy effects of macromutation were seen as a better fit to Kelvin’ belief about a young earth.
But by 1940 the “Modern Synthesis of Evolution” was established.
Growth of population genetics (using presigious mathematics) led to stress on the detrimental effect of most mutations, and the low probability of macromutations.
Gradualism returned assisted by experimental genetics. Dobzhansky simulated slow adaptations in fruit flies.

Wolf-Singer. The Brain: A Self-Organizing Learning System

1012 to 1014 neurons stretching 20,000 times around the equator.
Basic question: How can such a complex system develop in a safe and reproducible way from a comparatively small set of genetic instructions.

There is an interaction between genes and cellular environment, which determines what genes will be expressed. Microenvironment of the gene changes, changing gene expression, changing microenvironment, changing expression etc.
Also, electrical activity from developing sense organs provides an additional signalling system in the dialog.

This has important biological and untimately even philosophical implications.
A spiral of reciprocal interactions between the genome and its environment allows a rather small set of genetic rules to promote the development of systems as complex as the human brain.

Experience-dependent development: “Visual centers develop normally only when visual experience is available during a critical period of early development.” Patients who have suffered from congenital opacities of the eyes during early development and are therefore unable to perceive contours fail to recover visual functions after puberty when the optical media of the eyes are restored by surgery. The reason is that the genes alone are not capable of instructing the development of neuronal connections with sufficient precision.”
Binocular vision: Images from the two eyes must be precisely related in the brain. Brain selects correlated activity patterns from the parts of each eye that are getting the ‘same’ image -“activity-dependent selection.”

Experience-dependant pruning only occurs when both eyes are looking at the same unambiguous target. Arousal and attention systems, originating from the old central core of the brain, also participate.

The classical nature-nurture question of inherited versus acquired abilities can be re-addressed.

‘…The genome determines the general layout of neuronal connectivity and the basic response properties of neurons, many of which develop independently of experience.
These experience-independent specifications predetermine the criteria according to which the brain selects and categorizes sensory signals. Since patterns of sensory activation have to match these prespecified response properties to induce long term modifications, the genetic predisposition limits the range of stimuli that can influence self organization. The genes also determine the nature of possible relations that can be established by experience between the various feature domains. Associations can only be established between those sets of neurons that are interconnected to begin with. Furthermore the genome defines the rules according to which predetermined connections are selectively stabilized. We have seen that these modification rules have an associative function.” They provide “coherency” by reinforcing connections whenever two events occur simultaneously in space and time. (Remember the Triesman model) Gating systems determine whether a particular activation pattern should lead to long lasting modifications.

“Experience-dependent self-organization, therefore has little in common with passive instruction of a tabula rasa. Rather the developing brain appears to be a highly active and self-contained system that, at birth, already possesses substantial knowledge about the structure of the world to which it is going to adapt itself.” So at birth, the brain poses a number of precise questions to the environment to optimize and adapt its internal structure to reality. In a number of neuronal systems these questions are only raised during a brief and critical period.

“It soon becomes impossible to distinguish between cause and effect of certain developmental bifurcations.’
‘The developing brain and its environment appear as components of a highly interactive system whose complexity and richness far exceeds the information provided by the genome.
Philosophical question: Can consciousness and mind be considered as a property emerging from individual brains alone or must brains of similar complexity interact with one another during a long period of ontogenetic self-organization until mutual reflections between brains lead to the emergence of symbolic representations in the form of language and eventually to such concepts as those of self and consciousness.

Self-organizational coherency is like classical Pavlovian conditioning (learning).
Note on critical periods-imprinting: reduces confusion by confining learning to a period during which the probability is high that only members of the family will be around.
In view of the similarities between experience-dependant self-organization and learning it is not always possible to distinguish the two processes.

From A. R. Damasio: Descartes’ Error

Some Words on Neural Development. (pp. 108-113).

1. The human genome (the sum total of the genes in our chromosomes) does not specify the entire structure of the brain. There are not enough genes available to determine the precise structure and place of everything in our organisms, least of all in the brain, where billions of neurons form synaptic contacts. The disproportion is not subtle: we carry probably about 10 to the 5th (100,000) genes, but we have more than 10 to the fifteenth (10 trillion) synapses in our brains. Moreover, the genetically induced formation of tissues is assisted by interactions among cells, in which cell adhesion molecules and substrate adhesion molecules play an important role. What happens among cells, as development unfolds, actually controls, in part, the expression of the genes that regulate development in the first place. As far as one can tell, then, many structural specifics are determined by the genes, but another large number can be determined only by the activity of the living organism itself, as it develops, and continually changes throughout its life span.

2. The genome helps set the precise or nearly precise structure of a number of important circuits in the evolutionarily old sectors of the human brain, certainly the brain stem the hypothalamus and the basal forebrain, and probably the cingulate region and the amygdala. We share the essence of these brain sectors with individuals in numerous other species. The principle role of the structures in these sectors is to regulate basic life processes without recourse to mind and reason. The innate patterns of activity of the neurons in these circuits regulate homeostatic mechanisms without which there is no survival. Without them we would be unable to breathe, regulate our heartbeat, balance our metabolism, seek food and shelter, avoid predators and reproduce. They also intervene in the development and adult activity of the evolutionary modern structures of the brain.

3. The remaining brain circuits develop as a result of interaction with the physical environment and other individuals, using as a basis the evolutionarily old circuits already developed. Both the records of experiences and the responses to them, if they are to be adaptive, must be evaluated and shaped by a fundamental set of preferences of the organism that consider survival paramount. Note discussion of modulator neurons on p 111.

This account underscores the inadequacy of conceiving brain, behavior, and mind in terms of nature versus nurture or genes versus experience. Genes provide for one brain component with precise structure and another for which the precise structure is to be determined. But the to-be-determined structure can be achieved only under the influence of three elements: 1. The precise structure; 2. Individual activity and circumstances (in which the final say comes from the human and physical environment as well as from chance); and 3. Self-organizing pressures arising from the sheer complexity of the situation.

Oyama: Ontogeny and the central dogma: Do we need the concept of genetic programming in order to have an Evolutionary perspective.

Note: remember my perspective on Dawkins’s programming metaphor: Where does the crunchy texture of the biscuit come from.

Oyama considers this huge multivariate semantic world which, in one way or another makes nature more equal than nurture.
Faces of ubiquity.

Genetic programming as shades of preformationism. Its surrogates “proliferate wildly in the psychological and biological literature.

Note Tinbergen’s 4 questions.

The central dogma: Crick: one way flow of information from genes in ontogeny.
(Remember Stent’s question: Who reads the program?) Things are self-organizing
Part of the problem: lack of accord between developmentalists and evolutionists. Embryologists know there is more to it than genetic determination.
Ambiguity of terms such as biological, inherited, genetic and maturational permits blurring of Tinbergen’s whys.

Problem of dualism. Interaction not too helpful. Proponents mistake compromise and relabeling for conceptual resolution. P 52. “The problem of attributing some parts of the phenotype to the genes and some to the environment is that developmental processes and products are simply not partitionable in this way.”
P 54 “conventional dualism fails then, because it rests on an incoherent mix of ideas: there is no consistent way of distinguishing features that are programmed from features that are not.”

P 55. “The problem with this imperialistic version of developmental dualism is that it is vacuous: a genotype has just these developmental possibilities that it has (though who is to say what they are). (Chomsky tells us it is all the features, meaning whatever we find is genetically programmed.

P 62 Problem of developmental psychologists taking a set of givens.
More equal problem not solved by environmentalism.
Nature-nurture dichotomy will continue to dominate as long as we continue to speak of traits, programs or encoded potential as being transmitted.
Solution: Nature must be regarded as a product, not as predetermination. Developmental systems perspective.
Culture as developmental context.
Note problem of computer programming metaphor.

P 57. “Scholars and lay people alike continue to distinguish necessary inner essence from contingent outer appearance.”
Relevance to Marler’s “Innately guided learning”. “The theorists attempts to replace the innate-acquired distinction with closed and open programs or with inherited ranges of possible forms or innate genetically determined epigenetic rules merely blur the distinction between traditionally conceived nature and nurture when they should be questioning the very basis of the distinction.”
Note: meanings of innate: 1. Species-typical; 2. Predetermined; 3. Conferring survival advantage,; 4 Unlearned; 5 Having an evolutionary history; and independent of the environment.

“If evolution is construed as change in the constitution and distribution of developmental systems, the study of ontogeny is no longer a poor relative, to be lent evolutionary legitimacy by hook or by crook. Rather it becomes the very heart of evolutionary biology.
Me: You could look at going from favoring reduplication to favoring variegation in this way. Important notion. Nurture comes into conflict with nature phylogenetically rather than simply complementing it trivially. Semantically speaking modern nature a la Chomsky comes into conflict with old nature.

P 63. “When constructive interaction is seen to be fundamentally important for the formation (not just the support) of all feature, including “biological” ones then the role of the environment is not complementary to that of biology, but is constitutive of it in much the same way the genes are.”

P 70 Our inheritance does include culture, not as a second set of traits transmitted by an extragenetic conduit, but as aspects of the developmental context.”
P 71 “The concept of developmental systems gives a unified view of development while integrating it with evolution.”

“What is missing from most accounts is the synthetic processes of ontogenetic construction”. THAT’S WHAT WE WE WILL PROVIDE.
Note Hamburger, p 76 on why embryology was not incorporated into the neodarwinian synthesis.

Gottlieb: Normally occurring environmental and behavioral influences on gene activity: From central dogma to probabilisitic epigenesis.

Note cross-species action of the same gene Fig. 2.
Note Stent: “too many removes” p 794.
Me: Who programs the programmer.
Note regulative proteins acting on genes. Gene expression.
See the diagrammatic conceptions of the central dogma and probabilisitic epigenesis. Me: Deafness as the improbable.
Wasps, goats, (Slijper’s goatSheldon-type identical twins, and hens teeth.
“Genes express themselves appropriately only in responding to internally and externally generated stimulation.
Note Table 2. Normally occurring environmental and behavioral influences on gene activity.
“environmentally-provoked gene expression is thought to be required for long term memory” See Goelet et al.
Ontogeny of the nervous system: the earliest synaptic conncentions in the embryonic and fetal nervous system are created by spontaneous activity of nerve cells.
“the range of possible adaptive genetic responses to strange environmental conditions is truly astonishing” p 800

Plotkin: Plato’s Question.
“the nature-nurture question is the crucible within which psychology’s relation to evolutionary theory must be forged.

This is not because there is any simple, hence separation, of nature onto historical cause and nurture onto now and forever proximate causes. There isn’t such a mapping. Both nature and nurture are to be explained by both sets of causes. Conceptual epiphany: nurture has itself evolved. Nurture can only be fully understood in the light of historical cause. Nurture has nature. P 37

Darwin: Confused. Instincts & intelligence. Didn’t put intelligence in an evolutionary context, but, valuably, emphasized descent with modification of human intelligence.
MacDougall: Instincts. Others – labels as explanations. Weakness of the stance allowed easy demolition by behaviorists.

American ideologically based flight from elitist biologism to egalitarian cultural relativism.
Voice in wilderness: Kroeber: culture “not only is in nature, but is wholly part of nature.” P 47
Basic premise: Use of “innate” not sinful. “Present actually or potentially at birth and part caused by genetic information.”

Ethology: the scientific study of animal behavior.
Lorenz: Darwinism requires historical causality.
Solution to the N-N problem: innate school marms – learning by instinct. Lorenz understood nurture has nature but was stuck with nature being more equal than nurture.
Developmental solution: Oyama et al.
“Genes only convey information in so far as they are embedded within self-organizing developmental systems.

Note: Naturalness superceded in Shannon’s “snoopy”.
“Learning is constrained”. “genetic shaping of those structures in the brain that subserve learning and memory.
Innateness label descriptive. We still need an explanation in developmental terms. P 71.
Beach: The dichotomy between instinct and reason was never challenged by Darwinian theory. Separate determinism.

To See and Not to See: Oliver Sacks

William Harvey: “Nature is nowhere accustomed more openly to display her secret mysteries than in cases where she shows traces of her workings apart from the beaten path.” 9 108.

Such cases, hypothetical or real, had riveted the attention of philosophers and psychologists for hundreds of years.
Locke: After sight restoration you couldn’t tell which was the sphere & which was the cube. Likewise Berkeley.

So it proved. From Cheselden’s 13 year old boy, in 1728 “nearly all had experienced the most Lockean confusion and bewilderment. (Only those blind from birth or early childhood.)
Virgil, 53, virtually blind since early childhood, had cataract removed from right eye.
First report: “Virgil can see … entire office in tears.” Next day, problems.
History. Seeing problems from birth. Triple illness at 3 – virtually blind. Became another person – from spunky mischeivous to indolent, nonchalant, passive.
Vision recovered significantly but retinas gravely damaged, whether disease provoked or congenital. Cataracts developed at 6 & he went to a school for the blind. Passivity remained. Became a massage therapist.

On the first moment: There was light, there was movement, there was color, all mixed up, all meaningless, a blur. Surgeon said “Well?” & he realized that this chaos of light and shadow was a face. Gregory’s patient S.B. had virtually the same experience (accidentally blinded at infancy, corneal transplant in fifties.) Virgil’s retina and optic nerve were active but what he saw had no coherence. (If sight is lost late in life cataract removal reveals normal world almost instantly.)

He could see colors & movements, large objects & shapes Letter reading at 20/80. Central vision poor and trouble fixating. Retina moth eaten or piebald.
Mentally blind. Didn’t really look at faces. To see but not to decipher.
“One does not see or sense or perceive in isolation- perception is always linked to behavior and movement, to reaching out and exploring the world. It is insufficient to see. One must look as well. Von Senden: bandages removed at 5 didn’t look. John Hull: 5 years after becoming blind (at 50) he didn’t look at people.

Coming from the airport, he liked movement and big things. Moon larger than expected. (Gregory’s patient expected a wedge, not a crescent.) Big stuck airplane was a blimp. Birds made him jump. Got scared at supermarket. Everything ran toegether. Enjoyed uncluttered views but had no idea of size and perspective. “Sensation itself has no markers for size and distance; these have to be learned on the basis of experience.
First month of seeing had been predominantly positive.

Found walking scary, without touch, without his cane. Confused by his shadow, came to a stop, tripped or tried to step over it. Steps seemed a confusion, a flat surface of parallel and criscrossing lines. 5 weeks after surgery he felt more disabled than before the operation, but hoped all this would sort itself out with time. But “every patient in the literature had faced great difficulties after surgery in the apprehension of space and distance-for months and even years. (Valvo’s patient H.S. blinded at 15, corneal transplant at 22: “street lights were luminous stains stuck to the window panes, and the corridors of the hospital were black holes.)

At 5 weeks, confused his dog and cat. Amy: “You’d think once was enough.
Cheselden: at first learned to know and then to forget a thousand times a day.
Ability to do letters but not faces was apparently due to tactile experience and cross-modal transfer. But couldn’t read or see the words. H.S. had similar problems. Could not even count his own fingers. Virgil could pick up details but not synthesize them. “Trees did not look like anything on earth. A month after op: Virgil finally put a tree together- he now knows that a trunk and leaves go together to form a complete unit.” Skyscrapers strange, cannot understand how they stay up without collapsing. TG: I could no longer coordinate visual sensations with my speed of walking ( in getting to obstacles and an timing the appropriate response to them. Valvo: The real difficulty here is that simultaneous perception of objects is an unaccustomed way to those to sequential perception through touch. Sacks: the blind live in a world of time alone. The idea of space becomes incomprehensible – even for highly intelligent people blinded relatively late in life. (Von Senden’s main theme). Hull: Space is reduced to to one’s own body, and the position of the body is known not by what objects have been passed by but by how long it has been in motion. People are not there unless they speak. Hull after 5 years had to trace a three (tactile motor) to determine which way round it went. The recognition of color and movement seems to be innate. Seeing colors above all had a physical and emotional impact, almost shocking. Virgil distinguished colors but often gave them wrong names.

Virgil believed he had visual memories but were they just verbal like Helen Keller.
First he was unable to recognize any shapes, even a circle & a square which he recognized instantlly through touch.

Solid objects much more difficult cos their appearance was so variable. Much of the 5 weeks devoted to them: “their unexpected vissicitudes of appearance as they were seen from near or far, or half concealed, or from different places and angles.

His house was first unintelligible. He had to be shown around in detail – each chair.
Established canonical lines through the house. “It was only from this line, at first, that he could recognize anything-though this took a great deal of interpretation or inference.” A whiteness to the right was the dining room table. If he deviated from the line, he was totally disoriented. He started to make small investigative seeing-feeling sallies from the lines. Sacks was remined of an infant moving his hand to and fro. (Me: maybe an analog of babbling, or of Max throwing oven trays around.) Note Cezanne consciously moving his station point around – I could be occupied for months simply by bending to the left or right.
We achieve perceptual constancy – the correlation of all the different appearances, the transforms of objects in the first few months of life. It’s a huge learning task even the largest supercomputers cannot begin to match. For Virgil it required hours of conscious and systematic exploration of the smaller things each day- fruits, bottles, cans, cutlery, flowers, the knickknacks on the mantlepiece- turning them round and round, holding them close to him, and then at arm’s length, trying to synthesize their varying appearances into a sense of unitary objecthood. S.B. never ceased to be “struck by how objects changed their shape as he walked round them. He would look at a lamppost, walk round it and stand studying it from a different aspect, and wonder why it looked different and yet the same.”

All newly sighted subjects, indeed have radical difficulties with appearances, finding themselves suddenly plunged into a world that for them may be a chaos of continually shifting unstable evanescent appearances. They may find themselves completely lost in this flux of appearances. The newly sighted are baffled by the very concept of “appearance” which, being optical, has no analog in the other senses. (Note Bradley: Appearance and Reality (l893). After 5 weeks no difficulty with common objects, but unfamiliar objects were much more difficult. Flummoxed by blood pressure cuff until he could touch it. Moving objects presented a special problem for their appearance changed constantly. “Even his dog, he told me, looked so different at different times that he wondered if it was the same dog. Problems with faces and their expressions are almost universal. S.B. couldn’t recognize faces or their expressions a year after, despite perfectly normal elementary vision. Also no success at all on pictures. Virgil did not comprehend the idea of representation. (p 129.) Cheselden’s patient suddenly discovered pictures represented solid bodies but was then amazed that the objects were not 3-D to touch. He asked, which was the lying sense, feeling or seeing. Virgil, despite being an avid listener to baseball games could understand them at all on TV if the sound was turned off.

Virgil did some very crude drawings.
Gregory’s patient did not see illusions, (many seen in young children and some in monkeys) by parallel lines diverging, the Neckar cube as 3D and reversing, the wife and the grandmother. “..illustrates how rudimentary his brain’s powers of visual construction are.
At the zoo, he was surprised by an EMU’s movement, guessed it was a kangaroo from its hops, but could not describe it unless he could touch it. Probably identifed animals by motion or by a single feature, but he could not form any overall impression. Also problem of distinguishes objects (e.g. elephants) from background. He thought a gorilla looked human, but after fluently exploring a statue, He said “its not like a man at all”.
Note S.B. seeing and touching a lathe: “Now I’ve felt it, I can see.”
Virgil started to buy toy soldiers etc to capitalize on this seeing by feeling route.
Started eating well with knife and fork but gradually deteriorated and reverted to eating with his hands.

Note Virgil’s extremely sensitive auditory system: “They’re feeding the lions.” “Same with shaving.” Compare Sacks looking at EEGs.
But he also had periods of blurriness some lasting several days. Unique to the opthamologist.

(Normally half the cortex is given over to visual processing.)
Brain systems in all animals respond to overstimulation by shutdown.
It is not a world that one perceives or constructs but one’s own world. Note Anton’s syndrome, p 136. Perceptual self may collapse, altering the orientation and the very identity of the individual. If this occurs, an individual not only becomes blind but ceases to behave as a visual being, offers no report of any change of inner state, is completely oblivious of his own visuality or lack of it.

Wedding: Virgil’s family treated him as blind and he passively became blind. Not like Anton’s syndrome but more top-down or psychic. (Under stress asthmatics become more asthmatic, and parkinsons become more parkinsonian. Von Senden – motivational crisis.
S.B. went into depression and died 2 years after the operation.
Diderot; Problem of their blindness is ours not theirs.
Takeover of normally visual areas of the brain may be part of the problem of sustaining visuality after cataract removals.

Q. of whether the learning analogy should be with the newborn. Not really. Reorganization not organization. Thus the person must go from time to space.
Note on Hebb ( 140). Gregory: The patient is faced with reversing “the perceptual habits and strategies of a lifetime.” A Valvo patient: “One must die as a blind person to be born again as a seeing person.” It is the interim, the limbo—between two worlds, one dead / the other powerless to be born, that is so terrible.

Page 142. Essential note on analogy with problems of the deaf.
After right eye cararact removal”he was amazed at the range of skin colors he saw,and slightly disgusted by blemishes and “stains” in skins that to his hands had seemed perfectly smooth. S.B: “He also found some things he loved ugly (including his wife and himself!) and he was frequently upset by the blemishes and imperfections of the visible world.
Went to nutcracker, saw people jumping around but couldn’t see what they were wearing.
Xmas: great joy at seeing home and family.

Things seemed optimistic at Xmas but on Feb 8 he contracted pneumonia.
Showed blindsight –avoidance of obstacles he claimed he could not see. Sacks: visual cortex knocked out but subcortical centers remained.

Vision became virtually nonexistent. No electrical activity in retina or cortex.
One question: did overexposure to visual input burn out his retinas.
More likely: effects of chronic hypoxia.

At this stage he showed rage: “rage and his helplessness and sickness, rage at the smashing of a promise and a dream, and beneath this, most fundamental of all, … rage at being thrust into a battle he could neither renounce or win.”
Release was given in the form of the final blindness.

M.S. Gazzaniga, R.B. Ivry & G.R. Mangun, Cognitive Neuroscience, 2nd Ed.
Ch 3. pp. 70-94

GROSS AND FUNCTIONAL ANATOMY OF THE NERVOUS SYSTEM

Cerebral Cortex

Outer 1.5-4.5 mm “bark” of the hemispheres. Mostly cell bodies, hence gray. Much of the inside is fiber tracts of axons, hence white because of the myelin (Fig. 3.8)
Four lobes: Frontal, Parietal, Temporal, Occipital. (Fig. 3.9) Gyri & Sulci. (Fig. 3.7). (Central sulcus = Rolandic Fissure; Lateral sulcus = Sylvian Fissure. (Jump to Fig. 3.18 to see primary cortex (direct projection areas) secondary cortex and tertiary cortex)
Hemispheres joined by Corpus Callosum (a Commisure) (Fig. 3.8).
Finer divisions: Cytoarchitectonics – Brodmann’s areas (3.10).

Motor Areas of the Frontal Lobe. (Fig. 3.12) A. Precentral Gyrus – Motor Strip – Brodmann’s Area 4 is Primary Motor Cortex (M1). Direct output to Brain Stem and Spinal Cord. B. Anterior to this is Brodmann’s area 6. Two components: Premotor Cortex laterally, and the Supplementary Motor Area medially. These also have some direct output but also output to M1. C. Most anterior region is Prefrontal Cortex. “… takes part in the higher aspect of motor control and the planning and execution of behavior, tasks that require integration over time.” (p 49). Has 3 main areas: See Fig. 3.12.

Somatosensory Areas of the Parietal Lobe. Primary Somatosensory cortex (S1) is in the postcentral gyrus (Brodmann’s Areas 1, 2, 3.) (Fig. 3.13) Receives information from the thalamus on touch, pain, temperature sense and limb proprioception (position sense). S1 projects ventrally to S2. Both project to Posterior Parietal cortex.

Visual Processing Areas in the Occipital Lobe (fig. 3.14). Primary Visual Cortex – Striate cortex – V1 – Brodmann’s area 17: receives input from lateral geniculate nucleus of the thalamus.
Located medially. Codes visual features such as color, luminance, spatial frequency, orientation, and movement. Two pathways:
1. Retina, optic nerve, lateral geniculate, cortex.
2. Superior colliculus – cortex (visual-motor information).
Extrastriate cortex: Brodmann’s 18 and 19. More than three dozen subareas. (Fig. 3.16) Two major striate-to-extrastriate pathways (Fig. 3.15): 1. Ventral “What” pathway to Inferotemporal cortex, for form discrimination and object identification. 2. Dorsal “Where” pathway to Parietal cortex, for motion and visuospatial localization. Much but not total interconectivity between subareas including much reciprocity.

Auditory Processing Areas in the Temporal Lobe. Primary Auditory Cortex (AI) is Heschl’s Gyrus, in Superior Temporal Lobe inside the Sylvian Fissure (Fig. 3.17). AII is surrounding auditory cortex (Brodmann’s 41 & 42). A 22 surrounds this.
Tonotopic localization – reception layed out in terms of frequencies (in Hertz - Hz).

Association Cortex. This is cortex which is not sensory or motor. Receives input from more than one modality. Important for higher mental processes.

Limbic System, Basal Ganglia, Hippocampus, Hypothalamus and Diencephalon

Limbic System.
Limbic Lobe: (Fig. 3.19). Cingulate Gyrus, Parahippocampal Gyrus, Subcallosal Gyrus, Dentate Gyrus, & Hippocampal Formation. These structures interconnect with each other and with subcortical structures such as the Amygdala, Hypothalamus, Thalamus and Basal Ganglia to form the Limbic System (Fig. 3.20). Limbic lobe has primitive cortex (<6 layers) phylogenetically older than neocortex.
Limbic system participates primarily in emotional processing, but also in learning and memory.

Basal Ganglia.
A collection of subcortical neuronal groups in the anterior forebrain.
(Fig. 3.21) Caudate and Putamen = Neostriatum. With Globus Pallidus (Paleostriatum) they form the Corpus Striatum. With the Substantia Nigra and the Subthalamic Nucleus, they participate in background control of movement. Also involved in short term memory and executive functions. Major hookup: cortico-subcortical loop (Fig. 3.22) suggests monitoring function.

Hippocampal Formation and Medial Temporal Lobe.
Fig. 3.23. Medial temporal lobe and hippocampus involved in memory and amygdala involved in emotional processing.

Diencephalon
Thalamus and Hypothalamus. Thalamus: top of brain stem (see ventricular view) See Fig. 3.25. 2 thalami connected via massa intermedia (Fig. 3.23). “Gateway to the cortex”. Last precortical sensory way station.
Lateral Geniculate (Fig. 3.26): retino-cortical relay station.
Medial Geniculte: cochleo-cortical relay station.
VPL and VPM: Somatosensory relays.
Also in loops with basal ganglia, cerebellum and cortex.
Pulvinar connects with posterior cortex. Connections with parietal cortex for attentional control and with parts of the where-is-it system.
Cortical relay nuclei receive return cortical information via the thalamic reticular nucleus.
Hypothalamus (Fig. 3.27) Important for the autonomic nervous system (Sympathetic and Parasympathetic components; see 3.33) the endocrine system, maintenance of homeostasis, and for emotional processing. Releases peptide hormones influencing other neural sites.

Brainstem

3 parts: Mesencephalon (Midbrain); Metencephalon (Pons) and Myelencephalon (Medulla). (Fig. 3.28) Groups of sensory and motor nuclei, nuclei of widespread modulatory neurotransmitter systems, and white matter tracts of ascending sensory information and descending motor signals. Also controls respiration and sleep & wakefulness. Inner core: Reticular formation. Controls arousal and other functions.

Mesencephalon (Midbrain) (Fig. 3.29)
Superior Colliculus: Eye Movements. Visual reflex functions.
Inferior Colliculus: Auditory relay nucleus.

Pons & Medulla (Hindbrain) (Fig. 3.30)
Ascending sensory & descending motor pathways. Pons bulge related to pathways to cerebellum.

Cerebellum (Fig. 3,31)

Inputs to deep nuclei, and cortex, and outputs to deep nuclei. Inputs from sensory and motor centers, and vestibular input. Output to thalamus and thence to motor and premotor cortex. Projections to brainstem nuclei influence descending motor pathways to spinal cord.
The cerebellum is key to maintaining posture, walking and regulating coordinated movements.

Spinal Cord (Fig. 3.32)

Sensory (Dorsal and Motor Ventral) relay functions and simple reflex functions. Central grey matter like butterfly.