Bunge - From Neuron to Mind

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    NEWS IN PHYSIOLOGICAL SCIENCES

    Neuron to Mind

    main approaches to the study of mankind, and the corresponding

    of man, are sketched and examined. The systemic model of man

    a biopsychosocial entity is recommended. Likewise the multilevel

    is favored over the holistic, the analytic, and the synthetic

    Finally, the question of the reducibility of psychology

    neurophysiology is discussed.

    there was little com-

    between physiologists and

    ists. Typical ly the former were

    in the various systems and

    stems of the organism, whereas the

    studied behavior. Hardly anyone

    interested in the mechanisms where-

    neural systems control behavior,

    less in the nonmotor and nonsen-

    of the central nervous

    tem (CNS) of higher vertebrates. Con-

    mind nonexistent.

    That gap is now closing. Neuroscien-

    are becoming more interested in

    or, perception, emotion, and idea-

    while psychologists and ethologists

    happily ignoring the paralyzing in-

    “Do not neurologize ” and are

    to speculate on that which con-

    and does the mentation.

    fusion of neurophysiology with psy-

    ogy, which Karl Lashley demanded

    half a century ago, is fina lly start-

    to happen.

    Neurophysiologists are becoming

    re that the nervous system is only

    subsystem of the whole animal-

    most interesting of all-and

    hologists are realizing that real ani-

    not black boxes. The great wall

    body and behavior is being

    from within (starting with the neu-

    and from without (starting with be-

    r). As the boring proceeds we real-

    that the wall is not in nature but in

    and theology.

    Yet the fusion or merger tactics have

    far been delineated in vague terms.

    should introduce some precision if

    want to know how best to integrate

    Bunge is Professor of Philosophy in the

    and Philosophy of Scienc e Unit,

    University, 3479 Peel St., Montreal,

    I-DA 1 W7, C anada.

    the various approaches, methods, and

    results of the various sciences, from neu-

    rophysics to sociology, concerned with

    the problem of accounting for behavior

    and mentation. In the firs t place we

    should tackle the matter of the variety

    of approaches to this problem and weigh

    their comparative merits.

    Five approaches to the study

    of mankind

    Humans can be studied from different

    points of view: as a physical entity, as a

    chemical system, as an organism, as a

    psychosystem, and as a component of

    social systems (family, firm, school, etc.).

    Each of these approaches has its virtues

    and shortcomings, and each suggests a

    one-sided view of man.

    Approach 1 (physical). No doubt this

    is the basic approach and one that has

    proved fertile. But restricting the study

    of man to its physical components and

    aspects, i.e., adopting physicalism, is ig-

    noring whatever physics fails to explain.

    And trying to reduce the supraphysical

    features of man to physics is quixotic if

    only for being impractical. The mere at-

    tempt to write down, and even more to

    solve, the Schroedinger equation for a

    brain, or even a neuron or an amino acid

    molecule, is mind boggling. But even if

    such tasks were feasible, a number of

    essential features of life , such as evolu-

    tion, would remain in the dark. And

    overlooking evolution amounts to for-

    saking the understanding of the com-

    plexities of the nervous system and the

    mental and social peculiarities of hu-

    mans.

    Approach 2 (chemical). First the

    chemical theory o f heredity, then molec-

    ular biology, and finally neurochemistry

    showed that biochemistry is just as im-

    portant a tool as biophysics for under-

    standing the nervous system. However,

    the successes of the chemical approach

    should not be exaggerated, if for no other

    reason than that we stil l do not have an

    adequate understanding of a single or-

    ganic molecule. (To be sure chemists

    have discovered the composition and

    gross structure of millions of molecules,

    but they still owe us accurate theories of

    even the simplest of molecules involved

    in cell chemistry. Bear in mind that not

    even HZ0 has been calculated quantum

    mechanically.) In short, chemism is in-

    adequate.

    Approach 3 (biological). This is of

    course the approach characterizing the

    life sciences. This platitude bears repe-

    tition in view of the fact that the alter-

    native approaches are still going strong.

    (Thus most psychologists believe that

    they can afford to ignore biology, partic-

    ularly neuroscience and the theory of

    evolution.) And it also goes without say-

    ing that the biological approach can and

    should be combined with the other ap-

    proaches rather than being adopted to

    their exclusion. The latter stand, namely

    biologism, is one-sided for it tends to

    forget the physical and chemical com-

    ponents of living matter as well as the

    psychological and social dimensions of

    man.

    Approach 4 (psychological). The

    proper task of psychology is of course the

    understanding of behavior and ideation

    patterns. To be sure, all such functions

    are biological: they are performed by liv-

    ing organs; hence all good psychology is

    basically psychobiology. However, the

    functions of the nervous systems of com-

    plex organisms are different from those

    of simpler systems such as the cardio-

    vascular or the digestive ones. For ex-

    ample, the former can learn and per-

    ceive, two functions that are far beyond

    the capability of all but highly evolved

    organisms. Yet such behavioral and

    mental abilities should not lead to psy-

    chologism, i.e., the claim that psychology

    owes nothing to biology or to sociology,

    which is what behaviorists and psy-

    choanalysts have been saying.

    Approach 5 [sociological) . There is no

    understanding the social animal apart

    from social science. In particular, con-

    sciousness and language seem to be prod-

    ucts (and in turn modifiers) of social life ,

    and so are stress, moral conscience, and

    organizational ability. However, it would

    be a mistake to adopt sociologism, or the

    Volume 4/0ctober 1989

    0886- 17 14/4/89 150 0 1989 Irk Union Ph ysiol, Sci./A m. Physiol, Sot.

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    attempt to explain man in purely socio-

    logical terms, For, although sociali ty has

    iological and psychological roots, it

    transcends them. Indeed, unlike inver-

    tebrates and lower vertebrates, higher

    vertebrates possess social plasticity, i.e.,

    the capacity to readjust their patterns of

    social behavior in the face of internal or

    external difficulties.

    So much for the main legitimate, al-

    beit one-sided, approaches to the study

    of man. The engineering approach has

    not been included for the simple reason

    that animals are not machines: they are

    alive and have not been designed. To be

    sure, there are some similarities between

    man and machine, in particular the com-

    puter; if there were none we would not

    use them as labor-saving devices. How-

    ever, no analog is a substitute for the real

    thing, particularly when the analogies in

    question are so superficial that they

    overlook the speci fically physical , chem-

    ical, biological, psychological, and social

    properties of man, such as rational think-

    ing, planning, and abil ity to set up and

    dismantle social groups. In short, ma-

    chinism is inadequate.

    To sum up, there are fi ve legitimate

    and frui tful approaches to the study of

    man. But the adoption of any of them to

    the exclusion of the others, while una-

    voidable given the enormity of the task,

    should be regarded as only a temporary

    expedient. We should carry on work

    along each of the five approaches and

    should attempt to integrate them, fo r hu-

    mans are complex systems exhibiting all

    five aspects.

    Eight models of mankind

    The understanding of a thing begins

    and ends with some conceptual model of

    it. The model is better the more inclusive

    and accurate that it is. But even rough

    models can be used to guide-or block-

    research. Each of the five approaches

    examined in the last section has given

    rise to a set of models of man, In addition,

    technological and religious approaches

    have resulted in a certain model. Finally ,

    an eighth model is in the making, which

    brings together all five scienti fic ap-

    proaches.

    The first , or religious model, is that of

    Plato and Christian theology. According

    to it man is a spiritual being that uses

    his body as a tool during his temporary

    sojourn on earth. (As Sir John Eccles put

    it, the self is to the brain what the pianist

    is to the piano.) This animistic model

    was demolished by Darwin and by phys-

    iological psychology, but it still lingers

    on among philosophers, psychologists,

    and even neuroscientists unable to

    match their philosophy with their sci-

    ence. It has effect ively slowed down the

    merger of neurophysiology with psy-

    chology.

    The second, or technological model,

    regards man as a complex information

    processor, This model has captivated the

    imagination of many neuroscientists and

    psychologists . To be sure the neuroen-

    docrine system is, among other things,

    an information system; so is the neu-

    roendocrine-immune supersystem. How-

    ever, it happens to be a biosystem, i.e.,

    one characterized by biological proper-

    ties. And computer science has no room

    for such speci fic properties, not even for

    specifi c physical and chemical proper-

    ties. (As far as computer science is con-

    cerned an information system can be

    built out of modules of almost any kind.)

    Besides, it is a mistake to compare bio-

    systems to artifacts, for this suggests that

    the former, too, have been designed to

    some purpose. Finally , machinism is in-

    compatible with evolutionary biology,

    for machines are not subject to genetic

    mutation or natural selection.

    The third, or physicalist model, pre-

    supposes not only that physics is the

    basic science, which is true, but also that

    in principle no other science is neces-

    sary, which is false. Not even chemistry,

    the nearest neighbor, is reducible to

    physics without further ado. First, this is

    because chemical systems have peculiar

    properties such as lack of inertia that

    physics knows nothing about, and sec-

    ond, physical theory, although necessary

    to understand chemistry, is insuff icient:

    one must add subsidiary assumptions

    concerning, e.g., chemical composition

    and structure, that go beyond physics.

    The fourth or chemicalist model o f a

    human as a chemical reactor fares better

    than the physicalist model because i t

    is richer and, after all, the cell is con-

    stituted by chemical subsystems. But

    of course there is more to life than

    just chemistry. In particular, animals

    equipped with a neuroendocrine sys tem

    have neural and hormonal control sys-

    tems in addition to the genetic control

    system. To be sure all such control sys-

    tems are physicochemical, but they hap-

    pen to regulate biofunctions such as

    feeding, reproduction, self-repair, and

    defense.

    The fif th or biologistic model of man

    although far superior to the preceding

    ones, is defective in that it neglects the

    speci fically human abilities and short-

    comings. In particular, it overlooks the

    unique psychological and social features

    of human beings. Thus i t fosters the an-

    cient animistic doctrine of the soul as a

    separate entity. The practical conse-

    quence is clear, namely an unwilling-

    ness to use physical , chemical, and bio-

    logical means to modify behavior and

    mentation and the insistence on the po-

    tency of logotherapy.

    Likewise the sociologistic model of

    humans exaggerates one aspect at the

    expense of all others. This has two un-

    desirable consequences. One is to view

    society as a whole rather than as a sy

    tem and therefore as existing by itse

    and above the individual. The other con

    sequence is to deny that there are di

    orders of nervous cells and neural sy

    tems and blame all behavioral an

    mental disorders on society as a whole

    (This is of course the g ist of antipsychia-

    try. ) Like psychologism, sociologism i

    nores biology and thus is scient ifical ly

    unacceptable.

    Finally the eighth model, or system

    ism, pictures man as a biosystem com

    posed of numerous subsys tems, eac

    with its own specific functions, as we

    as a component of suprabiological (s

    cial) systems. This view includes what

    ever is valuable in previous models

    indeed the systemic model of humans

    acknowledges physical and chemical

    properties as well as biological, psycho

    logical, and social ones. In particular,

    man as a whole as well as his ever

    component possesses physical properties

    such as mass, but from the cell upward

    all the subsystems have peculiar supra

    physical properties, i.e., features n

    studied by physics. Also primates ca

    feel and dream, imagine and plan,

    well as enter into social relations an

    thus modi fy other animal’s mentation

    and behavior

    -all of which lie beyond

    physics and chemistry although rooted

    to the peculiar physical and chemical

    properties of living tissue.

    In conclusion, there are (at least) eigh

    models of man, or rather kinds of model

    Two of them, the religious and the e

    gineering models, are unscienti fic, an

    five are scienti fic but one-sided, for eac

    accounts for just one side of the whole

    Only the systemic model brings together

    whatever is valuable in each of the fiv

    one-sided or partial scientific models, b

    depicting man as a biopsychosocial sy

    tem with physical and chemical compo

    nents,

    Systems and levels

    Every concrete thing is either a system

    or a component of a system, i.e., a thin

    composed of interconnected things.

    particular, a human being is a compo

    nent of several social systems (family

    firm, club, etc.) and is in turn composed

    of a number of macrosystems, in partic

    ular the nervous system, which are

    turn composed of smaller subsystems

    This “hierarchical” organization goes

    to the world sys tem and down to the ce

    level and even further, to the level

    cellular subsystems (e.g.‘, ribosomes) an

    their molecular components (see Fig.

    This being so, to understand the behav

    ior of each module we must understand

    its components, environment, and struc

    ture, as well as the supersystem of which

    it is a component. One may despair

    the complexity of the task but may tak

    consolation in the fac t that the task is

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    performed by the entire scientific

    nity, perhaps with the help of

    eager to emphasize the

    for integrating the various partial

    Every leve l of organization is a set

    material things, each of

    ch is characterized by peculiar prop-

    particular laws). The interlevel

    ion in the “hierarchy” (or better

    el structure) is this. Any system be-

    to a given level is composed of

    to preceding levels. The

    ily of levels is thus ordered by the

    relat ion defined in this way.

    So far our sketch of the level structure

    been static. But we know from var-

    studies, in particular from studies

    development and on evolution, that

    leve l structure is far from being given

    In fact we know that

    stem has self assembled (or self

    from things on the preceding

    l (i.e., every system on a given leve l

    preceded in time by its components,

    ch are therefore rightly called its pre-

    The systems of any given level have

    properties in common with their

    and others that the latter

    are their emergent properties.

    an atom has an energy

    spectrum that its individual components

    do not possess; likewise, a molecule has

    an energy spectrum that is not the mere

    superposition of the spectra of its com-

    ponent atoms; and a neuronal system has

    a connectiv ity that is absent from its

    components. In short, at every level

    some properties (in particular laws) are

    gained (or emerge) while others are lost

    (or submerge). In short, there is both

    emergence and submergence along the

    evolutionary process.

    The methodolog ical morals of the pre-

    ceding ontologica l considerations are

    quite obvious:

    1) identify the level(s)

    crossed by your object(s) of study; 2) do

    not skip any levels; and 3) recognize the

    genealogy of the higher levels. These in-

    junctions help evaluate research strate-

    gies and projects. We may distinguish

    four main such strategies or methodol-

    ogies:

    Holism. Holism is the study of every

    thing as a whole and only on its own

    level, e.g., the study of the brain as a

    whole by means of the electroencepha-

    logram or the holographic model of

    memory.

    Analysis. Analysis is the reduction of

    the system to its components, or “top

    down.”

    Svnthesis. Synthesis is the bui lding up

    J

    World system

    (e.g. hypothalamus)

    (e.g. ribosomes)

    Elementary particles and fields

    The supersystems and subsystems of a human being.

    Social levels

    I

    Biological levels

    Chemical levels

    hysical levels

    of the whole from its components, or

    “bottom up.”

    Multilevel approach. The multilevel

    approach is the study of every system on

    its own level as well as a component of

    a supersystemand ascomposedof lower-

    level things.

    Each of the first three strategies has

    its virtues and shortcomings. Holism em-

    phasizes that the whole has emergent

    properties but refuses o explain them in

    terms of composition and structure, so t

    borders on irrationalism. The analytic

    method stresses he importance of the

    composition of a system but misses ts

    emergent properties. The synthetic

    method does not have the defects of the

    other two methods but it is not always

    practicable; for instance, so far biologists

    have not synthesized a living cell. I sub-

    mit that the multilevel approach is the

    best of all four, for it recommends study-

    ing each system on its own level as well

    as on its adjoining levels.

    The multilevel approach is an eclectic

    or catch-as-catch-can strategy, allowing

    one to use whatever approaches, tech-

    niques, models, and data that may seem

    promising. Hence it is integrative, al-

    though not holistic, an undeniable merit

    at a time when excessive specialization

    leads to artificial fragmentation. In par-

    ticular, it is the one strategy capable of

    bringing together all the studies in neu-

    roscience and psychology and thus the

    one capable of bridging the gap between

    neuron and mind.

    The multilevel strategy has proved its

    mettle in a number of domains. Thus the

    solid-state physicist builds a mathemat-

    ical model of his crystal structure the

    ion lattice together with the electron

    cloud) to explain such molar properties

    as electrical and thermal conductivity.

    The quantum chemist, even when intent

    on adopting a purist or ab initio syn-

    thetic or bottom up) approach, makesuse

    of whatever knowledge he can get from

    both classical chemistry and atomic

    physics. And the neurobiologist studying

    a particular system, such as the Wer-

    nicke “area,” approaches it on at least

    three levels: as a system of neurons, asa

    molar system with peculiar properties,

    and as the organ of the formation and

    understanding of linguistic expressions.

    Explaining behavior and mentation

    The most ancient and popular expla-

    nation of behavior is of course the an-

    imistic or mentalistic one: it takes the

    mind for granted and attributes to it the

    ability to control behavior. The barren-

    ness and untestability of mentalism

    drove psychologists to the denial of the

    mental, whereas the right attitude

    should have been to regard the mental

    assomething to be explained rather than

    as self-explanatory. Behavioristic psy-

    chology attempted to dispensewith the

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    and to describe behavior in terms

    stimuli. But, because it ignored the

    system, behaviorism gave a su-

    icial account of behavior-just as su-

    as the description of motion

    by pre-Newtonian kinematics.

    Biopsychology, on the other hand,

    admitting the valid results of be-

    goes far beyond it by looking

    (or neuroendocrine)

    t “mediate” (actually

    behavior. Thus it pro-

    centr ifugal ly, from the CNS to be-

    For example, it attempts to ex-

    voluntary movement in terms of

    specific activity of certain neuron

    es located in the frontal lobes.

    than regarding the CNS as a

    information processor restricted to

    (or encoding) external stim-

    physiological psychology has learned

    e CNS has a largely autonomous

    activity, that nerv-

    activity is modulated by environ-

    rather than being

    determined by them.

    Physiological psychology is not re-

    to the study of behavior: it also

    s mentation where it exists, from

    perception, and imagery to de-

    n and self-consciousness. The

    is the same in a ll cases, namely

    tackle the data of observation and self-

    as problems, to make neu-

    mechanisms of behavior and subjec-

    and to check such con-

    by means of further observa-

    measurements, or experiments.

    ate goal is of course to put such

    together into neatly stated

    (or special theories) of the var-

    behavioral and mental processes.

    would like to know the modus

    ndi of the smallest neuron assem-

    feeling fear or anxiety, of

    up a proposition, or of making

    Traditional (mentalistic) psychology

    s “pure” or untainted by physiology,

    it dealt with the putatively immater-

    oul or mind. Physiological psychol-

    the other hand, is based on (pre-

    poses) neurophysiology and, indeed,

    other branches of biology as well,

    h as endocrinology, immunology , and

    besides, it needs social

    nce to help explain some of the

    functions, such as moral con-

    Thus physiological psychology

    effect brings together al l the studies

    to the understanding of behav-

    and mentation. In other words, phys-

    gical psychology contributes power-

    y to the synthesis we are after, the

    capable of bridging the gap between

    d whole animal , between pro-

    sses at the subcellular leve l and bio-

    Is this synthesis a reduction, in partic-

    ular a reduction of psychology to neu-

    rophysiology? Not quite , and this for the

    following reasons. First, even at lower

    levels the derivation of one theory from

    another usually requires premises not

    contained in the reducing theory. Sec-

    ond, neuroscience itself needs the guid-

    ance of psychology; thus the study of

    perceptual systems is a matter not only

    for neurophysiology but also for the psy-

    chology of percept ion, which takes into

    account characteristics of the environ-

    ment, sometimes even of the social en-

    vironment. (Recall that perceptual error

    can be caused by social factors such as

    social pressures.) Third , there is more to

    neurobiology than neurophysiology,

    namely developmental and evolutionary

    biology. This poin t deserves clarifica-

    tion.

    Every contemporary organism is the

    outcome of two different processes: the

    result of an ontogenetic process and a

    product of a mult imill ion year evolu-

    tionary process. Either way nature ac-

    complishes the integration we find so

    difficult to conceptualize. Indeed the

    processes leading from molecule to fer-

    tilized ovum to adult primate, and from

    primitive cell to highly evolved animal,

    are processes of self-assembly (or self-

    organization), hence integrative.

    This has become a platitude, yet it is

    apt to be temporarily forgotten by the

    electrophysiologist or the psychologist.

    In fact, the division of scientific labor has

    reached such a ridiculous extreme that

    many workers in neuroscience and psy-

    chology tend to pay only lip service to

    the importance of studies in develop-

    ment and evolution for the understand-

    ing of their subject. Such neglect of

    development and evolution has had un-

    desirable consequences, such as 1)

    overlooking the biolog ical maturation of

    the CNS, which, in the case of certain

    systems such as the corpus callosum,

    takes up to a decade, and 2) exaggerating

    leaps at the expense of graduality (as is

    the case with mentalistic psychology,

    particularly of the information-process-

    ing variety, and its refusal to learn from

    animal psychology), or, conversely, 3)

    exaggerating continuity at the expense

    of quantitative novelty (as in the case of

    animal psychologists who claim that hu-

    man mental abilities differ only in de-

    gree from prehuman ones).

    To sum up, behavior and mentation

    can be explained with the help of neu-

    roscience and social science. The new

    psychology, based on neuroscience and

    willing to learn from social science, is at

    the very heart of the attempt to explain

    behavior and mentation in scientific

    terms. And it constitutes a synthesis or

    merger rather than a reduction, even

    though the behavioral and mental proc-

    esses are neurophysiological. (A philos-

    opher would say that this is a case of

    ontological reduction

    temological reduction

    without ful l epis-

    .

    1

    Conclusion

    Physiology and psychology share one

    ultimate goal, namely the scientific un-

    derstanding of behavior and mentation.

    No one branch of physiology or of psy

    chology can attain this goal single-hand-

    edly, because he problem is a multilevel

    one, and this becauseman himself exists

    on all levels. Nor should philosophy be

    left aside, because t too is interested in

    the mind-body problem and can offer

    valuable suggestions or facilitating the

    integration or synthesis of disciplines

    that the problem calls for.

    Neurophysics, neurochemistry, neu-

    robiology, psychology, and sociology

    may be pictured as forming a pentagon

    Philosophy lies at the center of it, some

    times blocking the whole view, as in

    psychophysical dualism; at other times

    favoring the integration of research

    fields; and at all times giving and receiv-

    ing stimuli from the other disciplines.

    To be sure one must specialize if one

    wants to make original contributions to

    knowledge. But specialization need not,

    nay must not, exclude the elaboration of

    a general scheme of things allowing one

    to locate one’s problems and thereby to

    make use of any relevant scraps o

    knowledge found in other fields. Such

    integration is indispensable in bridging

    the gap between neuron and mind.

    This paper is based on the author’s books

    A World of Systems [Boston, MA: Reidel,

    1979) The Mind-Body Problem [New York:

    Pergamon, 1980), Philosophy of Science and

    Techn ology, Part 2 (Boston, MA: Reidel, 1985),

    and Philosophy of Psychology, with Ruben

    Ardila [New York: Springer-Verlag, 1987), as

    well as on his article s “From neuron to behav-

    ior and mentation: an exercise in levelman-

    ship,” in Information Proce ssing in the Nerv-

    ous System, edited by H. M. Pinsker and W.

    D. Will iam s (New York: R aven, 1980, p. l-16),

    and “From mindless neuroscience and brain-

    les s psychology to neuropsychology” (Ann.

    Theor. Psych ol. 3: 115-133, 1985).

    I 1

    Volume 4/0ctober 1989 NIPS 20

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    Fifty Years Ago in Physiology

    In a real sense, 1989 has been the

    anniversary of modern cellular

    In 1939, J. Z. Young (3)

    d the full description, given

    shorter accounts two or three

    s earlier, of squid giant axons.

    lly arising from the fu-

    of smaller fibers, they were rec-

    as single nerve cells, a dem-

    revolutionized the

    . In that same year, building on

    of Gasser and Erlanger,

    o had developed the cathode ray

    for measuring electrical poten-

    nerves essentially free of time

    acts, Hodgkin and Huxley (2)

    st published together. They re-

    from inside a giant fiber im-

    ed with a Ag-AgCl-sea water

    and showed directly that

    potential is a transmem-

    event and that its magnitude

    overshoots the resting

    l (50 mV). And Cole and

    (1) succeeded in demonstrat-

    that action potentials in squid

    accompanied by a 40-fold

    in membrane impedance with

    change in capacitance, all new

    omena in 1939 for which the

    bases are now well established.

    Cole, K. S., and H. J. Curtis. Electrical

    impedance of the squid giant axon during

    act ivi ty. J. Gen. Physiol. 22: 649-670,1939.

    Hodgkin, A. L., and A. F. Huxley. Action

    potentials recorded from inside a nerve

    fibre. Nature Lond. 144: 710-711, 1939.

    Young, J. Z. Fused neurons and synaptic

    contacts in the giant nerve fibres o f ce-

    phalopods. Phil. Trans. R. Sot. Lond. B

    Ser. 229: 465-503,1939.

    Forty Years Ago in Physiology

    Almost every issue of every jour-

    reporting on cell physiology and

    biology contains at least one pa-

    in which the authors describe

    experiments. A com-

    n purpose is to determine trans-

    more and more experimen-

    are using micropipettes for in-

    mRNA into Xenopus oocytes

    express a desired protein product.

    inject fluorescent or other

    into a variety of cells to

    cell-cell communication,

    antibodies to block specific cyto-

    c functions, or foreign pro-

    measure their interactions

    h endogenous metabolites, or any

    a myriad of other compounds lim-

    NIPS

    Volume 4/0ctober 1989

    ited only by the experimenters’

    imaginations. All of this was made

    possible in 1949 by Ling and Gerard

    (l), who used micropipettes as elec-

    trodes to impale frog muscle cells.

    Others, including Ralph Gerard and

    colleagues, had attempted this be-

    fore but had found that the pipettes

    damaged the surface, resulting in lo-

    cal short-circuiting current leaks

    and, in consequence, disturbingly

    variable, unstable, and transitory re-

    sults. Ling and Gerard showed that,

    given sufficiently small tip diame-

    ters

    (Cl

    pm), the cell membrane

    sealed around the impaling pipette

    and permitted highly reproducible

    and stable (days) determinations of

    the membrane’s electrical proper-

    ties. What followed is history.

    1. Ling, G., and R. W. Gerard. The normal

    membrane potential of frog sartorius fi-

    bers. I. Cell. Comp. Physiol. 34: 383-396,

    1949.

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