neuros01.jpg (22089 bytes) brain.gif (2672 bytes) neuro001.gif (95862 bytes)
The psychological criteria for mind of George John Romanes (1848-1894) provide a starting point for modern studies. As a research associate of Charles Darwin he was one of the first to deal with mental functions in evolutionary terms.  

Known by his friends as "The Philosopher," he made fundamental contributions to neuromuscular physiology and evolution, as is set out in his original papers (reproduced in these web-pages) and in my books. On the other hand, he failed to understand the important messages Samuel Butler was attempting to convey and clung too intently to the Lamarckist aspect of Darwin's theory of pangenesis.

In the 1890s he established the Romanes Lectures at Oxford. The first lecturer was William Gladstone. The second was Thomas Huxley ("Darwin's bulldog"). The third was August Weismann whose doctrine of the continuity of the germ-plasm Romanes had initially questioned. The lectures continue to this day, supplemented by the Oxford London Lectures, which are also supported by Romanes Funds.

Donald Forsdyke


Romanes and Psychology (Contributions to psychology, metaphysics and language.)

Continuity between Mind of Man and Animals (Response to Sully's criticism (1891) of Romanes' book Mental Evolution in Man.)

Romanes & Neuromuscular Physiology (1944. Muscle fatigue on repeated electrical stimulation.)

Neural_Pacemakers (1965. Neuromuscular observations in a simple system - jellyfish. Homage to Romanes.)

Synapse_Concept (Anticipation of Sherrington by Schaefer)

Long Term Memory - Romanes and Samuel Butler (2008. Ideas that harken back to Avicenna.)

Long Term Memory - Scaling to Brain Size (2014. Evidence from adults who survived childhood hydrocephaly.)

Theory of Mind. Romanes and Butler (2015. Philosophical aspects - information ideas in biology - selfish gene history.)

Neuronal Plasticity and Spatial Learning (Human lymphocyte genes also affect brain function.)

Schizophrenia (Articles by Patricia Forsdyke, 1998, 2009)

Romanes and Psychology

Supporting Darwin, Romanes wished to establish a continuity between the evolution of the brains of animals and of man, so combating the "Rubicon" view promulgated by Max Muller and Alfred Wallace. To this end he studied various cognitive functions of animals and human children, but Ethel Romanes put her foot down when he suggested that a monkey he was observing should be allowed to interact with their children in the nursery. 

Animal Intelligence 1882

Mental Evolution in Animals 1884 (with a posthumous essay on instinct by Charles Darwin)

Mental Evolution in Man 1888 (A copy of this was among the books salvaged from the library of Sigmund Freud after he left Vienna, and has markings in the margin in Freud's hand. For Romanes' response to a critic: Click Here )

Excerpt from Romanes'  Animal Intelligence (1888 edition)

Now, in this mode of procedure what is the kind of activities which may be regarded as indicative of mind? I certainly do not so regard the flowing of a river or the blowing of the wind. Why?

  • First, because the objects are too remote in kind from my own organism to admit of it; and,
  • secondly, because the activities which they present are of invariably the same kind under the same circumstances; they afford no evidence of feeling or purpose.

In other words, two conditions require to be satisfied before we even begin to imagine that observable activities are indicative of mind:

  • first, the activities must be displayed by a living organism; and
  • secondly, they must be of a kind to suggest the presence of two elements which we recognize as the distinctive characteristics of mind as such—consciousness and choice.

So far, then, the case seems simple enough. Wherever we see a living organism apparently exerting intentional choice, we might infer that it is conscious choice, and therefore that the organism has a mind. But further reflection shows us that this is just what we cannot do; for although it is true that there is no mind without the power of conscious choice, it is not true that all apparent choice is due to mind. In our own organisms, for instance, we find a great many adaptive movements performed without choice or even consciousness coming into play at all—such, for instance, as in the beating of our hearts. And not only so, but physiological experiments and pathological lesions prove that in our own and in other organisms the mechanism of the nervous system is sufficient, without the intervention of consciousness, to produce muscular movements of a highly co-ordinate and apparently intentional character.

Thus, for instance, if a nervous connection between his brain and lower extremities, on pinching or tickling his feet they are drawn suddenly away from the irritation, although the man is quite unconscious of the adaptive movement of his muscles; the lower nerve-centres of the spinal cord are competent to bring about this movement of adaptive response without requiring to be directed by the brain. This non-mental operation of the lower nerve-centres in the production of apparently intentional movement is called Reflex Action, and the cases of its occurrence, even within the limits of our own organism, are literally numberless. Therefore, in view of such non-mental nervous adjustment, leading to movements which are only in appearance intentional, it clearly becomes a matter of great difficulty to say in the case of the lower animals whether any action which appears to indicate intelligent choice is not really action of the reflex kind.

On this whole subject of mind-like and yet not truly mental action I shall have much to say in my subsequent treatise, where I shall be concerned among other things with tracing the probable genesis of mind from non-mental antecedents. But here it is sufficient merely to make this general statement of the fact, that even within the experience supplied by our own organisms, adaptive movements of a highly complex, and therefore apparently purposive character, may be performed without any real purpose, or even consciousness of their performance.

It thus becomes evident that before we can predicate the bare existence of mind in the lower animals, we need some yet more definite criterion of mind than that which is supplied by the adaptive actions of a living organism, howsoever apparently intentional such actions may be. Such a criterion I have now to lay down, and I think it is one that is as practically adequate as it is theoretically legitimate.

Objectively considered, the only distinction between adaptive movements due to reflex action and adaptive movements due to mental perception, consists in the former depending on inherited mechanisms within the nervous system being so constructed as to effect particular adaptive movements in response to particular stimulations, while the latter are independent of any such inherited adjustment of special mechanisms to the exigencies of special circumstances.

Reflex actions under the influence of their appropriate stimuli may be compared to the actions of a machine under the manipulations of an operator; when certain springs of actions are touched by certain stimuli, the whole machine is thrown into appropriate movement; there is no room for choice, there is no room for uncertainty; but as surely as any of their inherited mechanisms are affected by the stimulus with reference to which it has been constructed to act, so surely will it act in precisely the same way as it always has acted.

But the case with conscious mental adjustment is quite different. For, without at present going into the question concerning the relation of body and mind, or waiting to ask whether cases of mental adjustment are not really quite as mechanical in the sense of being the necessary result or correlative of a chain of physical sequences due to a physical stimulation, it is enough to point to the variable and incalculable character of mental adjustments as distinguished from the constant and foreseeable character of reflex adjustments. All, in fact, that in an objective sense we can mean by a mental adjustment is an adjustment of a kind that has not been definitely fixed by heredity as the only adjustment possible in the given circumstances of stimulation. For were there no alternative of adjustment, the case, in an animal at least, would be indistinguishable from one of reflex action.

It is, then, adaptive action by a living organism in cases where the inherited machinery of the nervous system does not furnish data for our prevision of what the adaptive action must necessarily be—it is only here that we recognize the objective evidence of mind. The criterion of mind therefore, which I propose, and to which I shall adhere throughout the present volume, is as follows:--

Does the organism learn to make new adjustments, or to modify old ones, in accordance with the results of its own individual experience? If it does so, the fact cannot be due merely to reflex action in the sense above described, for it is impossible that heredity can have provided in advance for innovations upon, or alterations of, its machinery during the lifetime of a particular individual.

colorbar.gif (2362 bytes)

Romanes and Metaphysics

Romanes was led to psychology as part of a life-long quest for meaning in the universe. At Cambridge (1873) he won the Burney prize, awarded by Christ's College, for his essay on "Christian Prayer considered in relation to the belief that the Almighty governs the world by general laws". This was published in 1874 with an appendix on "The Physical Efficacy of Prayer". As the anonymous "Physicus", he then wrote "A Candid Examination of Theism" (1878), where the influence of Darwinian ideas is very strong, and the conclusion is most poignant:

Creation of life on earth (left) and the stars (right) from the Museum at Salerno in Italy.

View of divine creation. At left God creates the earth and at the right God creates the stars. From the Museum at Salerno, Italy.

"So far as I am individually concerned, the result of this analysis has been to show that, whether I regard the problem of Theism on the lower plane of strictly relative probability, or on the higher plane of purely formal considerations, it equally becomes my obvious duty to stifle all belief of the kind which I conceive to be the noblest, and to discipline my intellect with regard to this matter into an attitude of the purest scepticism.

    And forasmuch as I am far from being able to agree with those who affirm that the twilight doctrine of the "new faith" is a desirable substitute for the waning splendour of "the old", I am not ashamed to confess that with this virtual negation of God the universe to me has lost its soul of loveliness; and although from henceforth the precept to "work while it is day" will doubtless gain an intensified force from the terribly intensified meaning of the words that "the night cometh when no man can work", yet when at times I think, as think at times I must, of the appalling contrast between the hallowed glory of that creed which once was mine, and the lonely mystery of existence as now I find it, -- at such times I shall ever feel it impossible to avoid the sharpest pang of which my nature is susceptible.

     For whether it be due to my intelligence not being sufficiently advanced to meet the requirements of the age, or whether it be due to the memory of those sacred associations which to me at least were the sweetest that life has given, I cannot but feel that for me, and for others who think as I do, there is a dreadful truth in those words of Hamilton, -- Philosophy having become a meditation, not merely of death, but of annihilation, the precept know thyself has become transformed into the terrific oracle to Oedipus --

  "Mayest thou ne'er know the truth of what thou art.""

In 1878 Romanes was strongly tested by Thomas Huxley when asked to become a member of The Association of Liberal Thinkers, an Association designed to attack all forms of humbug in Victorian life, particularly religious "humbug." Romanes was not so sure that the time was right for discarding what Karl Marx had called "the opium of the masses." Romanes declined to join the Association in a letter dated Jan 3rd 1879. Two days later he sent a second letter explaining his views more fully, and concluding:

"Of course I have no doubt that in the long run Truth, however hideous, must prevail!"

In 1879 he married the deeply religious Ethel Duncan. Thus began an even stronger testing!

In his incomplete and posthumous "Thoughts on Religion" (1896) there was an attempt to return, through reason, to his previous position. Some of this was expressed in poetry, aspects of which has been extensively analyzed (Pleins, 2014). His eldest child "Fritz" (Ethel Georgina) was 14 when Romanes died of a brain tumour in 1894. She studied theology at Oxford and became a nun (see "The Story of an English Sister" by her mother (Romanes, 1918).

    On coming down from Cambridge, Romanes had worked in London with Professor John Burden Sanderson, whose later move to the Physiology Department at Oxford paved the way for Romanes' move there in 1890. Here he and Ethel Romanes entertained Burden Sanderson's nephew (John Scott Haldane) and his wife (Louisa Kathleen Trotter), who became in 1892 the mother of John Burden Sanderson Haldane (the "JBS" who gave us "Haldane's Rule" in 1922). 

"Jack" (John Burdon Sanderson Haldane) circa 1900

Louisa K. Trotter (Haldane) (1863-196?), friend of George J. Romanes and mother of J. B. S. Haldane. Picture circa 1890.

Louisa Haldane provides the following memory in her "Friends and Kindred" (1961):

"The two houses which I enjoyed going to were the Dicey's and the Romanes'. My first dinner party was at the Romanes', and he was quite definately Romanes of Nigg, not an unfamiliar Oxford professor.

     I saw a good deal of him during his last illness. The excuse was that he liked to hear about some experimental work of John's and we talked at large about people and things, and on the last few visits rather tentatively about religion. He spoke very restrainedly of the many visits he had from the clergy his wife believed in. 'They mean well,' he said, but they tired him terribly, and would try to make him answer questions.

     On my last visit, he held on to my hands and said, 'Don't go, don't go,' but what can one do when a man's devoted wife almost pushes one out of the room? I am quite confident that his death-bed confession was merely the result of being too exhausted to argue. It was not meant as acquiescence to Anglican doctrines."

As for the quest for meaning, the remarks of L. C. Dunn at the beginning of his A Short History of Genetics (1965) must suffice:

"If the time-depth is shallow, so, too, in the restricted study I have in mind will be the degree of philosophical penetration. The results achieved in the development of genetics were derived from the application of experimental methods, but the question which experiment will not answer is what it all means. 

    The hope for an answer to such a question, whether expressed or lurking in the background, may be an actual handicap to investigation. We may find ourselves unready or unwilling to accept the limited, either-or kind of decision which experiment is competent to provide and thus lose the solid substance of discovery for the shadow of a larger hope. 

    It is not that I, or another who may review the development of the science of heredity, may not be interested in what it means, but rather that before we ask that ultimate question we should know what has caused it to be asked."

colorbar.gif (2362 bytes)

Romanes and Language

David J. Murray (Queen's University at Kingston) has described Romanes' great contributions to our understanding of how human language evolved. Some recent articles:

  • Language and Psychology: 19th Century Developments Outside Germany: A Survey (2001). History of the Language Sciences: An International Handbook on the Evolution of the Study of Language. Edited by S. Auroux, K. Koerner, H.-J. Niederehe and K. Versteegh. Walter de Gruyter, Berlin, pp. 1679-1692.

  • Romanes (1888) on the Origins of Human Language. Proceedings of Workshop on The Unexploited Legacy. (2006) Max-Planck Institute for Psycholinguistics, The Netherlands. This includes a brief summary of Romanes' life.

Further assessments of Romanes may be found in:

  • Anctil, M. (2015) The experimental approach to jellyfish neurobiology. George Romanes and Theodor Eimer. In: Dawn of the Neuron: The Early Struggles to Trace the Origin of Nervous Systems. McGill-Queen's Univ. Press. Chapter 5.

  • Boakes, R. (1984) From Darwin to Behaviourism: Psychology and the Minds of Animals. Cambridge Univ. Press. Chapter 2.

  • Knoll, E. (1986) The science of language and the evolution of mind: Max Muller's quarrel with Darwinism. Journal of the History of Behavioural Sciences 22, 3-22.

  • Richards, R. J. (1987) Darwinism and the Demands of Metaphysics and Religion: Romanes, Mivart and Morgan. In: Darwin and the Emergence of Evolutionary Theories of Mind and Behavior. University of Chicago Press.

  • Murray, D. J. (1988) A History of Western Psychology. 2nd Edition. Englewood Cliffs, NJ. pp. 255-271.

  • Howe, M. L. & Courage, M. L. (1993) On resolving the enigma of infantile amnesia. Psychological Bulletin 113, 305-326, 305-326.

  • Searleman, A. & Herrmann, D. (1994)  Memory from a Broader Perspective. McGraw Hill, New York.

  • Murray, D.J. (1999) A "presence/absence hypothesis" concerning hippocampal function. [Commentary on J. P. Aggleton & M. W. Brown: Episodic memory, amnesia, and the hippocampal-anterior thalamic axis] Behavioral and Brain Sciences 22, 462-463.

  • Murray, D. J., Kilgour, A. R. & Wasylkiw, L. (2000) Conflicts and missed signals in psychanalysis, behaviourism, and Gestalt psychology. American Psychologist 55, 422-426. , 422-426.

  • Pleins, J. D. (2014) In Praise of Darwin. George Romanes and the Evolution of a Darwinian Believer. Bloomsbury, New York.

colorbar.gif (2362 bytes)

Neuromuscular Physiology

With the guidance of John Burdon Sanderson, Romanes studied neuromuscular physiology in the Jodrell Laboratory at University College, London. In May 1876 there was a reading at the Royal Society of his paper "Observations on the galvanic excitation of nerve and muscle, with special reference to the modification of the excitability of motor nerves produced by injury." (Journal of Anatomy & Physiology 10, 709-734). His pioneering observations were recognized by Ralston in 1944:

G. J. Romanes on the Excitability of Muscle

by H. J. Ralston (1944) Science 100, 123-124, 123-124.
College of Dentistry and Medicine, University of California

All students of evolutionary theory are familiar with the fundamental contributions of George John Romanes in that field, but it is to be regretted that his physiological studies are not nearly so well known. Inasmuch as the elucidation of the electrical and chemical factors underlying muscle fatigue is a most important objective of research in neuromuscular physiology, Romanes' work along this line should be recalled.

    In a letter to Charles Darwin dated August 13th 1877, Romanes says:

I am very glad you have drawn to my attention prominently to the localizing function in Drosera, as it is very likely I have been too keen in my scent after nerves; and I believe it is chiefly by comparing lines of work that in such novel phenomena truth is to be got at.

    And this reminds me of an observation which I think ought to be made on some of the excitable plants. It is a fact not generally known, even to professed physiologists, that if you pass a constant current through an excised muscle two or three times successively in the same direction, the responses to make and break become much more feeble than at first, so that unless you begin with a strong current for the first of the series, you have to strengthen it for the third or fourth of the series in order to procure a contraction.

    But on now reversing the direction of the current, the muscle is tremendously excitable for the first stimulation, less so for the second, and so on. Now this rapidly exhausting effect of passing the current successively in the same direction, and the wonderful effect of reversing it, point, I believe, to something very fundamental in the constitution of muscular tissue. The complementary effects in question are quite as decided in the jelly-fish as in the frog's muscle; so I think it would be very interesting to try the experiment on the contractile tissues of plants.

    The discovery of the above reported phenomenon is generally credited to Gulacsy, who reported it in 1929. When one considers that Romanes also first observed, in the umbrella of the jelly fish, what is now known as fibrillation, it would seem appropriate to take cognizance of his important physiological research by attaching his name to one of these phenomena. It is therefore suggested that the recovery of excitability of tissue upon reversal of polarity of a stimulating current be called the "Romanes effect".

colorbar.gif (2362 bytes)


Neural Pacemakers

 The basis of our understanding of pacemaker physiology, now seen as so important in heart research, was laid by Romanes in the 1880s. His appointments to the Linnaean Society and the Royal Society indicate that his contributions were recognized by his peers. He later won high praise when his results were confirmed using elaborate twentieth century technologies (Passano 1965; Satterlie 2002)

Pacemakers and Activity Patterns om Medusae: Homage to Romanes

by L. M. Passano (1965) American Zoologist 5, 465-481.
Department of Zoology, Madison, University of Wisconsin

Even though they were carried out over 90 years ago, there is still a marvelous freshness about the discoveries of G. J. Romanes concerning what we would now call the behavioral physiology of coelenterates. When periodically rereading his three classic papers in the Philosophical Transactions (1876, 1877, 1880), or his popular book (1885) culled from those papers, I am reminded again of his achievement: indeed a disquieting thought (which other investigators may have experienced also) is that Romanes has already observed and recorded all the facts of the matter, and that as our knowledge of these animals continues to increase, it only sustains his original insights. The more I learn of jellyfish, the more I understand what Romanes has to tell us of them.

    One of the features of this outstanding pioneer work was that Romanes used both hydrozoan and schphozoan medusae in his investigations, and drew general conclusions applicable to both. Since his day little physiological work has been done on the former assemblage of animals, so that with a few exceptions current knowledge is where he left it. Contrarily, a number of investigators have utilized the larger, less fragile scyphomedusae in the belief, first expressed by Romanes, that they should be simpler to understand as well as easier to work with. I, too, commenced working with scyphomedusae and have only recently been audacious enough to study various hydromedusae. That caution was justified; the latter are indeed more complex in their neurophysiology and behavior, so that my current work confirms Romanes' findings, but the hydromedusae are also much simpler to study with current electrophysiological techniques ... .

colorbar.gif (2362 bytes)

Sherrington's Synapse Concept

Following Romanes, a later student in Michael Foster's Cambridge School of Physiology was C. S. Sherrington (1857-1952). In 1970, it was pointed out that Romanes physiological studies on nerve conduction in jellyfish had inspired histological studies by his friend Edward Schaefer (later known as Sharpey-Schafer), which led to Sherrington's concept of the neurological synapse.


by R. D. French (1970) Medical History 14, 154-165.

French quoted the (1885) treatise - "Jelly-Fish, Star-Fish and Sea Urchins" - where Romanes summarized his work in invertebrate neurophysiology. In this book, after describing Schaefer's observations of anatomically discrete nerve fibres underlying the muscle of the jellyfish bell, Romanes anticipates Sherrington by writing:

"Now, if it is a remarkable fact that in a fully differentiated nervous network the constituent fibres are not improbably capable of vicarious action to almost any extent, much more remarkable does this fact become when we find that no two of these constituent nerve-fibres are histologically continuous with one another. Indeed it seems to me we have here a fact as startling as it is novel.

     There can scarcely be any doubt that some influence is communicated from stimulated fibre a to the adjacent fibre b at the point where these fibres come into close apposition. But what the nature of the process may be whereby a disturbance in the excitable protoplasm of a sets up a sympathetic disturbance in the anatomically separate protoplasm of b, supposing it to be really such - this is a question concerning which it would as yet be premature to speculate.

     But I think it may be well for physiologists to keep awake to the fact that a process of this kind probably takes place in the case of these nerve-fibres. For it thus becomes a possibility which ought not to be overlooked, that in the fibres of the spinal cord, and in ganglia generally, where histologists have hitherto been unable to trace any anatomical or structural continuity between cells and fibres, which must nevertheless be supposed to possess physiological or functional continuity - it thus becomes a possibility that in these cases no such anatomical continuity exists, but that the physiological continuity is maintained by some such process of physiological induction as probably takes place among the nerve fibres of Aurelia."

See also Satterlie RA (2002) "Neuronal control of swimming in jellyfish: a comparative story. Canadian J. Zoology 80, 1654-1669.

In a review in Endeavour (2013) "Cajal, Golgi, Nansen, Schaefer and the Neuron Doctrine" Ortwin Bock writes:

"The honour of being first to state the discontinuity of nerve cells goes to the Assistant Professor of Physiology at University College London, Edward Schaefer (1850-1935) who, as Edward Sharpey-Schafer, would later make many contributions to medical science. In a text received on 31 October 1877 and communicated to members of the Royal Society by his professor William Sharpey (1802-1880) on 10 January1878, Schaefer began: "Last August I undertook, at the request of my friend Mr. G. J. Romanes, an investigation with the view of proving the presence or absence of histologically differentiated nervous structures in the Medusae". He had used chloride of gold to stain the tissue and found that "If we trace out the course of the individual nerve fibres more closely . . .we are struck with certain remarkable facts . . . each fibre is entirely distinct from and nowhere structurally continuous with . . . any other fiber".


Neuronal Plasticity

While we can study, say, the liver, of experimental animals and obtain results which bear on human liver function, this is less likely when studying the brain. As Romanes argued, the human brain shares many features with those of animals. But the brain is special; it is the organ, par excellence, defining the human animal. If a giraffe were having problems with its neck, we could study the human neck instead, but it would be a poor substitute for studying the giraffe neck.

   Cells of different tissues use similar signalling mechanisms. Sometimes it is easier to study one tissue rather than another. A gene recognized in 1985 as playing a role in the activation of human lymphocytes, G0S30 (now commonly referred to as EGR1 or ZIF268), is involved in synaptic activation, spatial learning, and "reconsolidation" after memory recall, centred on the hippocampus. Similarly, G0S8/RGS2 (Click Here) is now recognized as playing a major role in neuronal plasticity and in synaptic development in hippocampal CA1 neurons. Its inactivation predisposes to anxiety and male aggression. G0S3/FOSB when ablated causes behavioural problems in mice. Inactivation of G0S9 (now known as NAMPT) leads to motor neuron dysfunction.

   Thus, targeting lymphocyte function, rather than brain function, produced results relevant to both tissues.  This human tissue, lymphoid tissue, can be readily obtained, live and functioning, from a sample of blood from a normal or sick person. It is not so easy to obtain live and functioning human neural tissue for research! Thus enforcing (through funding priorities) strict relevance to brain function may have delayed progress in brain research. (Click Here)

Fordyce, D. E. et al. (1994) Genetic and activity-dependent regulation of ZIF268 expression: association with spatial learning. Hippocampus 4, 559-568.

Ingi, T., Krumins, A.M., Chidiac, P., Brothers, G.M., Chung, S., Snow, B.E., Barnes, C.A., Lanahan, A.A., Siderovski, D.P., Ross, E.M., Gilman, A.G. and Worley, P.F. (1998). Dynamic regulation of RGS2 suggests a novel mechanism in G-protein signaling and neuronal plasticity. J. Neurosci., 18, 7178-7188. (Click Here for more on RGS proteins)

Lee, J. L. C. et al., (2004). Independent cellular processes for hippocampal memory consolidation and reconsolidation. Science, 304, 839-843.

Oliveira-dos-Santos, A. J., Matsumoto, G., Snow, B.E., Bai, D., Houston, F.P., Whishaw, I.Q., Mariathasan, S., Sasaki, T., Wakeham, A., Ohashi, P.S., Roder, J.C., Barnes, C.A., Siderovski, D.P. & Penninger, J.M. (2000). Regulation of T cell activation, anxiety, and male aggression by RGS2. Proc. Natl. Acad. Sci. USA. 97, 12272-12277.

Wang et al. (2017) Deletion of NAMPT in projection neurons of adult mice leads to motor dysfunction, neurodegeneration and death. Cell Reports 20, 2184-2200.

Wei et al. (2000) Role of EGR1 in hippocampal synaptic enhancement induced by tetanic stimulation and amputation. J. Cell Biol. 149, 1325-1333.

Worley, P,F. et al. (1991) Constitutive expression of ZIF268 in neocortex is regulated by synaptic activity. Proc. Natl. Acad. Sci. USA 88, 5106-5110.

Worley, P. F. et al. (1993) Thresholds for synaptic activation of transcription factors in hippocampus: correlation with long-term enhancement. J. Neurosci. 13, 4776-4786.

Romanes' Contributions to Psychology (Tuft's University) (Click Here)

Romanes' Evolution Studies, Correspondence, Genealogy, and Queen's University (Click Here)

colorbar.gif (2362 bytes)


Schizophrenia: Breaking the Silence (Click Here) traflite.gif (995 bytes)

colorbar.gif (2362 bytes)

Return to: HomePage (Click Here)

This page was established in 1999 and last updated on 03 September 2017 by Donald Forsdyke.

WWW page access counterSince 1st September 1999