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William H. Calvin and Derek Bickerton, Lingua ex Machina: Reconciling Darwin and Chomsky with the human brain (MIT Press, 2000), end matter.  See also

copyright ©2000 by William H. Calvin and Derek Bickerton

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This 'tree' is really a pyramidal neuron of cerebral cortex.  The axon exiting at bottom goes long distances, eventually splitting up into 10,000 small branchlets to make synapses with other brain cells.
William H. Calvin

University of Washington
Seattle WA 98195-1800 USA


We wish to thank the Rockefeller Foundation for hosting us for a month at their Bellagio Study and Conference Center at the Villa Serbelloni. We also benefitted from workshops organized by the La Jolla Origins of Humans group (sponsored by the Preuss Foundation and the Mathers Foundation) and the Center for Human Evolution at the Foundation for the Future. We got a lot of useful questions and advice from Yvonne Bickerton, Katherine Graubard, Ruth and Elihu Katz, and the other temporary residents of Bellagio; Jess Tauber, Peter " Throwing Words" Rockas, Elizabeth F. Loftus, Beatrice Bruteau, Blanche Graubard, Dan Downs, Chris Westbury, David Schoppik, Bart de Boer, Francis Steen, Gerhard Luhn, Heidi Lyn, Robert Berwick, Steven Pinker, Michael Rutter, and countless anonymous reviewers. We also thank John Sunsten, Stewart Brand, William Hopkins, Terry Deacon, Frans de Waal, Richard Dawkins, and Greg Ransome for helping us chase down illustrations and citations.



The bold face terms are cross-references to other glossary items; the all-caps items are the roles in argument structure. Many technical linguistic terms will not be found here, e.g., for "empty categories," consult the index and the appendix.

There is a more extensive neuro glossary in The Cerebral Code, available at The present glossary is also on the web at


agent A role (the performer of an action) in argument structure (JOHN cooked dinner).

allele Alternative versions of a gene. Perhaps 20 percent of your expressed genes have a different allele on the other chromosome; that is, you are heterozygous for that gene and might switch to using it under some conditions. One reason that hybrids donít breed true to type is that parents are often passing on their less-used allele. Inbred strains have less heterozygosity.

altruism Doing something for anotherís benefit, at expense to yourself Ė but not necessarily at the expense of your genes, as when you aid relatives. In reciprocal altruism (RA), sharing with nonrelatives is eventually reciprocated, though the system is weakened by freeloaders ("cheaters").

area When capitalized, itís a Brodmann Area, a subdivision of cerebral cortex based on the relative thickness of the six layers. Area 17 is better known as primary visual cortex; it seems to be a functional unit, but Area 19 comprises at least six major functional units. A territory or work space is a generic lower-case area, occupied temporarily by active patterns of cloned hexagons.

argument structure The assignment of thematic roles to the constituents (noun-phrases, prepositional phrases, even clauses) that represent the participants in actions, states, and events creates arguments. Some of these arguments are obligatory, depending on the meaning of the verb. Argument structure is what determines whether a verb will have one, two or three obligatory arguments. Thus, "sleep" will have only one, a sleeper. "Break" will have two, a breaker and something broken. "Give" will have three, a giver, a gift, and a receiver. Unlike phrase structure, argument structure does not assign either linear ordering or hierarchical relationships to its components; argument structure has to be mapped onto phrase structure in order to provide these.

axon The neuronís tail-like "wire," a long (0.1 - 2,000 mm), spider thread thin portion of the neuron that carries voltages between the neuronís input sites (concentrated on cell body and dendritic tree) and the neuronís outputs, its many-branched axon terminals that make synapses onto downstream neurons. Itís typically a one-way street, messages flowing from the dendrites and cell body to the far end of the axon where synapses are made.

bee dance The honeybee appears, at least in the context of a simple coordinate system, to have broken out of the usual animal communication that has a single meaning. When the bee returns to her hive, she performs a " waggle dance" in a figure_8 that communicates information about the location of a food source that she has just visited. The angle of the double racetrackís common axis communicates the direction of the new-found food, and the number of circuits that she does around the loops communicates the distance from the hive. But, as Bickerton said in Language and Species,

All other creatures can communicate only about things that have evolutionary significance for them, but human beings can communicate about anything.... Animal calls and signs are structurally holistic [and] cannot be broken down into component parts, as language can.... Though in themselves the sounds of [human] language are meaningless, they can be recombined in different ways to yield thousands of words, each distinct in meaning.... In just the same way, a finite stock of words... can be combined to produce an infinite number of sentences. Nothing remotely like this is found in animal communication.

beneficiary A role (doing something for someone) in argument structure (I bought it FOR YOU).

binding Binding theory describes the conditions that identify constituents without independent reference Ė pronouns and anaphors (reflexives, reciprocals, and the like). See the appendix.

bonobo Pan paniscus, formerly called the " pygmy chimpanzee," and the last great ape species to be identified. Until 1927, they were confused with the chimpanzee, Pan troglodytes Ė with the old common name arising because they were said to be the "chimpanzee of the pygmies," living as they do on the left bank of the Congo River in equatorial forests (theyíre called the "Left Bank Chimps" for other reasons, too). While of a more slender body build than chimps, bonobos are not particularly smaller. Humans last shared a common ancestor with both Pan species about 5 million years ago. The two Pan species diverged at about the beginning of the ice ages, 2.5 million years ago, roughly the same time that the Australopithecus lineage spun off the Homo lineage.

bottleneck In population biology, an occasion when genetic variability in a population was greatly simplified by the loss of alternative alleles Ė not because of selection against them but simply due to dropouts that occur by chance, because of reduced choice in mates. This happens when population numbers are greatly reduced for a time; the re-expansion then populates the world from the smaller range of variation.

central nervous system (CNS) The brain, spinal cord, and the retina (all the rest is the peripheral nervous system).

cerebral cortex The outer 2 mm (thatís two thin coins worth) of the brainís cerebral hemispheres with a layered structure. It isnít required for performing a lot of simple actions but seems essential for creating new episodic memories, the fancier associations, and many new movement programs. Paleocortex, and archicortex such as hippocampus, has a simpler structure and earlier evolutionary appearance than the six-plus-layered neocortex.

cheater detection A feature of reciprocal altruism theories, what you use to identify freeloaders who receive but rarely give back.

chunking Collapsing multiple-word phrases into a single word, in the manner of acronyms.

clause A verb and all its associated arguments. See phrase.

code In cryptography, a code is a disguising transformation that (unlike a cipher) also chunks Ė and thereby shortens Ė the message, as when a number stands for a standard five-word phrase. More generally, as in genetic code, code refers to the transformation of a representationís short form into its long-form implementation. As such, it is analogous to a matrix. Code may also simply refer to the short form itself, such as a DNA base-pair sequence capable of generating a particular protein.

columns A minicolumn is a cylindrical group of about 100 neurons about 0.023 mm in diameter, extending through all the layers of neocortex and usually organized around a dendritic bundle; the orientation column is an example. Macrocolumns are a hundred times larger in area (and about 0.4 -1.0 mm across) and often more like curtain folds than cylinders; they are typically identified by common inputs, e.g., the ocular dominance columns of visual cortex.

corticocortical connection An axon or axon bundle connecting one patch of cerebral cortex to another. Some remain local, within the superficial layers of cortex, while others go tunneling through the white matter to distant targets Ė some, via the corpus callosum, to the other cerebral hemisphere.

creole Children invent a new language Ė a creole Ė out of the words of the pidgin protolanguage they hear their immigrant parents speaking. A pidgin is what traders, tourists, and "guest workers" (and, in the old days, slaves) use to communicate when they donít share a real language; pidgin sentences are unstructured and short, while those in creoles have the features of universal grammar.

Darwin Machine The Calvin 1987 coinage, on the Turing Machine analogy, for any full-fledged Darwinian process incorporating the six essentials for the Darwinian algorithm. Species evolution, the immune response, some genetic algorithms, and the hexagonal work space competitions are all examples. Not to be confused with particular models, also named after Darwin.

dendrite Neurons have branches. At least in neocortex, dendrites are the receiving branches of the neuron and the axon is the sending branch. Elsewhere, some dendrites can also act like axon terminals, releasing neurotransmitter in response to impulses and local voltage changes. There is always a single thin axon that initiates and propagates impulses to distant destinations, and there are somewhat thicker dendritic branches that receive synapses from other neuronsís axon terminals. Pyramidal neurons have a tall tree-like apical dendrite plus some rootlike basal dendrites.

epigenetic rules Some aspects of development are both innate and specified by the environment; "epigenetic" applies to the environmental bits. For example, a plant has two growth states, a positively phototaxic mode that directs most growth toward the light, and a negatively phototaxic mode. Some vines initially grow away from the light; then, when there is enough root structure, further growth is up towards the light; the vine climbs any tree trunk and its branches. Then some vine tips turn negatively phototaxic, dropping a branch down to earth Ė which, upon reaching soil, digs in to create another set of roots. The behavioral switch may be provided by genes, but the overall form of the resulting plant also depends upon what was encountered in the environment along the way. Syntax too is likely to have epigenetic aspects; the sign-language-deprived deaf infants might be like plants with only oblique light.

exaptation This was a term intended to cover cases where an organ with one original function got adapted to perform another function (such as the swim-bladders becoming lungs when the first marine creatures became terrestrial). Previously, the term "pre-adaptation" had been used, but this form is objectionable insofar if it suggests some degree of prescience. (WHC: I use it anyway.)

gene A unit of heredity, essentially that segment of a DNA molecule comprising the code for a particular peptide or protein. We also talk loosely of "a gene for blue eyes" and so forth ( reification strikes again), but many a DNA gene is pleiotropic: it has multiple (and sometimes very different) effects on its body; like that maxim about intervening in complex systems, "You canít do just one thing."

genotype The full set of genes carried by an individual, whether expressed or silent alleles. Similar to genome. Compare to phenotype. What makes living matter so different from other self-organizing systems is that a cell has an information center, the genes, concerned with orchestrating the many different processes going on within the cell, and in such a manner that copies of the cell tend to survive.

goal A role (whoever the action was directed toward) in argument structure (I gave it TO MARY).

grammar Not to be confused with socially correct usage. In order to handle novel sentences, we not only need to access the words stored in our brains but also the patterns of sentences possible in a particular language. These patterns describe not just patterns of words but also patterns of patterns. There are three aspects of grammar: morphology (word forms and endings), syntax (from the Greek "to arrange together" Ė the ordering of words into clauses and sentences), and phonology (speech sounds and their arrangements). A complete collection of rules is called the mental grammar of the language, or grammar for short.

grammar, universal Each of the languages of the world has a corresponding mental grammar, constructed as we learn the language. Though they differ in many ways, the human brain seems to have a highly specific menu of possibilities for grammatical organization, known as Universal Grammar, or UG, that structures language learning even when the input itself is lacking in structure ( pidgins, home sign, and so on).

grammatical morphemes The words (a few dozen in English) that refer to relations between content words. They are unlike content words, which refer to concepts of things in the world. They include words that express relative location (above, below, in, on, at, by, next to), relative direction (to, from, through, left, right, up, down), relative time (before, after, while, and the various indicators of tense), and relative number (many, few, some, the _s of plurality). The articles express a presumed familiarity or unfamiliarity (the for things the speaker thinks the hearer will recognize, a or an for things the speaker thinks the hearer won't recognize) in a manner somewhat like pronouns. Others express relative possibility (can, may, might), relative contingency (unless, although, until, because), possession (of, the possessive version of _s, have), agency (by), purpose (for), necessity (must, have to), obligation (should, ought to), existence (be), nonexistence (no, none, not, un_), and so forth. These are called " closed-class words" because our ways of expressing relationships are so resistant to augmentation, whereas you can always create new nouns or verbs.

head See phrase.

impulse Action potential and spike are synonyms; an impulse is the regenerative change in the voltage across the neuronís membrane used for long-distance (more than a millimeter) signaling in the nervous system. It is brief (1/1000 sec, quicker than any other signal in the brain but a million times slower than computers) and large (only 1/10 volt but bigger than any other voltage in the brain). Its threshold property can also be used as a simple decision making mechanism. See also axon, myelin.

inflections The inflectional system of English alters a noun when it refers to a multiplicity ("The boy ate three cookie." Is that correct English?) and alters a verb when it refers to past time ("Yesterday the girl pet a dog." OK?). Late learners of English may fail to realize that anything is "wrong" with these incorrect sentences, as such long-range dependencies are redundant information that helps out in noisy environments when some words are imperfectly heard and must be guessed.

innateness "Hardwired by genes" is the general idea, but it is, more generally, a bit of behavior that arises without learning. When the individual finds itself in a particular setting, out pops some complicated behavior. Mating behaviors are innate; some things are too important to be left to learning. But there isnít a dichotomy between innateness and environmental causation; as epigenetic rules show, something innate may have, via environmental triggers, profound effects on future form and function.

instrument A role (doing it with something) in argument structure (Bill cut it WITH A KNIFE).

irregular An irregular noun does not follow the usual plural rule. By the age of two or three, children learn to add -s. Before that, they treat all nouns as irregular. But even if they had been saying "mice," once they learn the plural rule they may begin saying "mouses" instead. Eventually they learn to treat the irregular nouns as special cases, exceptions to the usual rule. Verbs too can be irregular: the past tense of "fly" is "flew" Ė unless, of course, the word is being used in some novel manner, as in "The batter flied out," where the regular -ed form is nonetheless used. Young children, however, often use the regular form for the central meaning of "fly," as in "The bird flied away."

inheritance principle Darwinís great but often misunderstood insight, that variation is not truly random. Rather than variations being done from some ideal or average type, small undirected variations are preferentially done from the more successful individuals of the current generation, exploring the solution space nearby (not jumping randomly to somewhere truly unrelated) in the next generation. []

island biogeography The peculiarities of animal and plant species when largely isolated, with just occasional interbreeding. An "island" can also be a deep ocean basin, a high mountain valley, or a patch in a patchy resource distribution that prevents migration. Islands often have a reduced number of species, so traditional predators or parasites may be lacking. Species often arrive in small numbers, so bottlenecks are a standard feature of island populations.

language acquisition device The LAD is a hypothetical mechanism in the human brain that enables any normal human to learn any of the 5000+ human languages (or any possible human language). Its existence has never been empirically demonstrated and, within the present approach, it would seem to be unnecessary to assume any LAD as a distinct, self-encapsulated unit.

meme Richard Dawkinsís 1976 coinage, on the analogy to gene (with a little aid from mime and mimic), for a cultural copying unit, such as the word or melody that is mimicked by others.

memory, dual trace Hebbís 1949 coinage for separate systems implementing short- and long-term memories: active (spatiotemporal) and passive (spatial-only) memory traces.

memory, episodic One-trial learning involving distinct episodes, such as being an eyewitness to an accident. Such memories are notoriously malleable, influenced by subsequent events and the mistakes made in earlier recall attempts.

movement Where a word moves out of its usual place in a sentence, as in the wh- words. See the appendix.

myelin The fatty, porcelain-colored insulation around an axon that reduces its energy consumption while also making the impulse travel much faster. It is wrapped in layers, like a bandage.


neocortex All of cerebral cortex except for such things as hippocampus, the simpler layered structure that lacks the patterned recurrent excitatory connections and columnar structures that make the six-layered neocortex so interesting.

nervous system The whole works, both central nervous system (CNS: brain, spinal cord, and retinas) and peripheral nervous system (most sensory and muscle connections, plus the clusters of neurons called ganglia).

neuron The nerve cell, whether sensory neuron, interneuron, or motor neuron. There are about 1012 neurons in the human brain and spinal cord; the neocortex alone is said to have 1011. The cell body of the neuron is the widest section, thanks to containing the cell nucleus, and there are many processes branching off, receiving inputs and distributing outputs. See dendrite, axon.

neurotransmitter A molecule such as glutamate or acetylcholine that is released from an axon terminal (often by the arrival of an impulse), diffuses across a narrow extracellular space, and binds with a receptor on the surface of the postsynaptic cell. (These three parts are collectively called the synapse). Many dozens of neurotransmitters have been identified over the years, and a given axon terminal may release more than one kind.

niche The "outward projection of the needs of an organism" such as food resources, migration routes, camouflage from predators, suitable housing and sites for effective reproduction. An empty niche is a proven niche space that is temporarily unoccupied by a tenant species.

parcellation Fragmentation; breaking apart a population into smaller, isolated units ("parcels" or "patches"), as when rising sea level converts a hilly island into an archipelago of former hilltops. See also island biogeography.

patient A role (who- or whatever undergoes the action) in argument structure (John cooked DINNER). Often described as theme.

phenotype Usually "body" but actually the entire constitution of an individual (anatomical, physiological, behavioral) resulting from the interaction of the genes with the environment. As Dawkins emphasized in The Extended Phenotype, can even include such things such as bird nests.

phoneme The units of vocalization distinguished by native speakers of a language. Unlike ape calls and cries, phonemes are all meaningless by themselves, having meaning only in combinations ( words). It is important to realize that phonemes are categories that standardize. For example, Japanese has a phoneme that is in between the English /L/ and /R/ in sound space. Those English phonemes are mistakenly treated by Japanese speakers as mere variants on the Japanese phoneme. Because of this " capture" by the familiar category, those Japanese speakers who can't hear the difference will also pronounce them the same, as in the familiar rice-lice confusion.

phrase A group of words consisting of a head (which can be a noun, verb, preposition, etc.) and its modifiers. Clauses consist of groups of phrases. Each phrase is labeled according to its head. If the head is a noun, the phrase will be a noun phrase ("the tall blond man," where "man" is the head). If the head is a preposition, the phrase will be a prepositional phrase ("with one black shoe," where the head is "with").

phrase structure The results of the procedure by which words and phrases are assembled to form clauses and sentences. Formerly an independent module in generative grammar, its features now fall out from the principles that govern other modules. However, phrase structure trees are still drawn by syntacticians to show the relationships between words, phrases, and clauses. Such trees are hierarchically structured and, nowadays, usually binary-branching.

pidgin A contact medium liable to spring up wherever speakers of several different languages have to communicate without any language in common. In its early stages of development a pidgin is a form of protolanguage: that is to say, it lacks any kind of formal structure. Pidgin utterances consist of small groups of content words strung together in a purely ad hoc fashion. A pidgin, if it endures long enough, may stabilize, expand and, after several generations, approach the status of a full natural language. If a pidgin, regardless of its stage of development, is acquired by children, they convert it into a creole.

postsynaptic The postsynaptic neuronís dendrite receives neurotransmitter, rather in the manner of sniffing perfume, and changes the permeability of its membrane to certain ions, usually Na+, K+, Cl-, or Ca++ in some combination. Ions flowing through the membrane in turn produce the voltage change known as the postsynaptic potential (PSP). If excitatory, it is called the EPSP; if inhibitory, the IPSP.

protolanguage Any form of communication that contains arbitrary, meaningful symbols but lacks any kind of syntactic structure. Forms of protolanguage include the communication of "linguistically"-trained apes and other animals, pidgins in their early stages, the speech of nonproficient second-language learners, and that of children under two.

pyramidal neurons The excitatory neurons of neocortex. They typically have a tall apical dendrite (an exception is the spiny stellate neuron) and a triangular-shaped cell body (from whence the name), from which their axon leaves. The neurons contributing to the pyramidal tract (alias the corticospinal tract, named for the triangular shape of the axon bundle as it traverses the medulla) are themselves pyramidal neurons, but most pyramidal neurons send axons elsewhere.

receptive field A map of the inputs to a single neuron, e.g., those parts of the skin of the hand that produce excitation or inhibition of a cortical neuron (antagonistic surrounds are especially common). The limited view of the world as seen by a single neuron. []

recombination There are several connotations: (1) The shuffling of genetic material between an individualís two chromosome pairs that occurs just prior to the production of ova or sperm (the crossing-over phase of meiosis); and (2) the production of a new individual through the union of a sperm and an ovum from two parents at fertilization.

schema As in "schematic outline," itís a mental item more abstract than a rich mental image of an object. In some cognitive contexts, it is used more narrowly for those things like more, less, bigger, inside Ė things grounded in our everyday experiences, often making reference to our own body moving through our daily world. Movements need something similar, and schema is often used to refer to standard movement programs.

semantics The "meaning" of words, those connotations that you might look up in a dictionary (as opposed to syntax).

source A role (taking it from someone or something) in argument structure (I bought it FROM FRED).

synapse The synapse is the junction between neurons across which communications flow, usually in the form of neurotransmitter molecules secreted by the presynaptic axon terminal that diffuse a short distance across the extracellular space (the synaptic cleft) to the postsynaptic neuron, on whose membrane are some receptor molecules to which the neurotransmitter molecules reversibly bind. While they are bound, they open up an ion channel through the postsynaptic membrane, producing postsynaptic current flow. Most drugs affecting the CNS operate by interfering with synaptic transmission. See also dendrite, neuromodulator, neurotransmitter, postsynaptic.

syntax The set of rules and principles that determine how sentences are formed, and the structures resulting from sentence formation.

theme A role (who- or whatever undergoes the action) in argument structure (John cooked DINNER).

UG See grammar, universal.

word order A simple convention that aids in identifying roles, such as the subject-verb-object order (SVO) of most declarative sentences in English ("The dog bit the boy") or the SOV of Japanese. At least in English, the who-what-where-when-why-how questions deviate from basic word order: "What did John give to Betty?" is the usual convention (except on quiz shows in which questions mimic the basic word order and use emphasis instead: "John gave what to Betty?"). Some languages such as Latin lack a systematic word order, instead using characteristic inflections or even separate words (as when English uses "he" for a subject and "him" for an object, although both have singular, masculine, third-person referents) to help disambiguate the sentence.


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Linguistics Appendix (DB)
About the Authors

Web supplement:  Some photographs of Bellagio.

Copyright ©2000 by
William H. Calvin and Derek Bickerton

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