Heredity as Transmission of


Butlerian Intelligent Design


Centaurus (2006) 48, 133-148

Author Posting. Copyright: The Author 2006. This is the author's version of the work. It is posted here for personal use, not for redistribution. The definitive version (with some imposed editing) was published by Blackwell Publishing Ltd. in Centaurus, 48:3, pp. 133-148.

Self-portrait (1878) of Samuel Butler (1835-1902). Reproduced with the permission of the Master and Fellows of St. John's College, Cambridge

Samuel Butler (1835-1902) 

With permission of the Master and Fellows of St. John's College, Cambridge.


1. Pococurante 

2. Samuel Butler in New Zealand

3. Samuel Butler in London

4. Heredity Equals Memory 

5. Memory as Stored Information

6. Variation

7. Lamarckism

8. Recent Controversy over "Intelligent Design" 

9. Conclusions

End Note on Dreams (Jan 2007)

End Note on Pernicious Anaemia (Apr 2010)

End Note (July 2010) - Commentary by Shaw



Abstract. In the 1870s Ewald Hering and Samuel Butler provided what was, for that time, a scientifically coherent foundation for the Lamarckist view that positive adaptations to the environment acquired during an individual lifetime can be transmitted to offspring. Observing that heredity was a form of memory (involving stored information), they distinguished what are now known as genotype and phenotype, and proposed that cognitive abilities present in the most elementary organisms might mediate a transmission of acquired adaptations. While compatible with the then available facts of evolution, this Butlerian version of "intelligent design" was rendered less credible by subsequent appreciations of the discrete (discontinuous) inheritance of many phenotypic characters (Mendelism), and of the separation of germ-line from soma (Weismannism). However, it can now be seen that twenty-first century bioinformatics has nineteenth century roots.

Keywords.  bioinformatics, Butler, Darwin, evolution, heredity, Hering, information, instinct, intelligent design, Lamarck, memory, Mendel, Romanes, variation, Wada

1. Pococurante

Slowly scanning the assembly before him, his stern gaze settled on one upturned face: "Pococurante!" Decades later Charles Darwin (1867) recalled the event: "I was at school at Shrewsbury under a great scholar, Dr. Butler; I learned absolutely nothing, except by amusing myself by reading and experimenting in chemistry. Dr. Butler somehow found this out, and publicly sneered at me before the whole school for such gross waste of time; I remember he called me Pococurante [a person concerned only with trifles], which, not understanding, I thought was a dreadful name."


2. Samuel Butler in New Zealand       

On Saturday October 1st 1859 Darwin (1809-1882) noted in his diary that he had finished correcting the proofs of his great work, The Origin of Species, which appeared in November (Bowlby 1990). That same Saturday, Dr. Butler's grandson, Samuel (1835-1902), sailed for New Zealand with Justus Liebig's Agricultural Chemistry and the aim of becoming a sheep farmer (Jones 1919, p. 74). Shortly after his arrival he obtained a copy of Darwin's book. Gregor Mendel is also likely to have obtained a copy in the early 1860s (Iltis 1932; Orel 1984). Thus, while Mendel was breeding peas and studying Darwin in Moravia, Samuel Butler was breeding sheep and studying Darwin in New Zealand. This inspired his first articles on evolution, which appeared in The Press of Christchurch and soon received the commendation of Darwin himself (Butler 1914a, pp. 149-154). The sheep breeding was financially successful. In 1864 Butler returned to England where he remained the rest of his life, becoming, in the words of one contemporary, "the most brilliant, and by far the most interesting of Darwin 's opponents." (Bateson 1909, p. 88).

    Here I review the concept of heredity as the transmission of stored information (memory) as introduced in the 1870s by Ewald Hering (1834-1918) and Samuel Butler. In the sense that the work of Mendel was quantally different from those of his intellectual ancestors but remains close to those of his intellectual posterity (Roberts 1929), so the work of Hering and Butler can now be seen as at the root of modern evolutionary bioinformatics (Forsdyke 2006). However, whereas Mendel's work escaped recognition for a "mere" thirty-five years, that of Hering and Butler has escaped modern commentators (Smith 1999; Smith 2000; Kay 2000) and lain hidden for over a century.


3. Samuel Butler in London

The man who gained posthumous prominence in the twentieth century as the author of Erewhon and The Way of All Flesh, was a grandson of Bishop Samuel Butler who was for many years the headmaster of Shrewsbury School in England. The Bishop's son Thomas and Charles Darwin were acquainted. Both attended the school and Cambridge University. Canon Thomas Butler said of Darwin that "he inoculated me with a taste for botany" (Jones, 1919, p. 12), but in 1854 he sent his son Samuel to Cambridge hoping he would enter the Church. Having studied mathematics and classics, Samuel graduated in 1858. Much to the consternation of his father, he declined to be ordained (Jones 1919, p. 61).

     Butler lived a short walk from what was then probably the greatest library in the world - that associated with the Reading Room of the British Museum. With a small assured income and, other than an annual visit to Italy, relatively inexpensive tastes, Butler had - what was denied the professional "men of science" whose writings he devoured - time. However, unlike the men of science, whom he came to challenge, his interests were not focused. He was attracted to painting and literature through which he hoped he might further his financial independence.

    In 1870 Butler collected together some earlier publications (Butler 1914a) with titles such as "Darwin Among the Machines," "The World of the Unborn," "The Musical Banks" and "An Erewhonian Trial," and wrote his most "successful" literary work, a satire on the materialism, pseudo-idealism and academic pretentiousness of his times - Erewhon (the title is close to "nowhere" backwards). It was completed in 1871 and went on sale in 1872 with the support of a loan to cover the publisher's expenses (Jones 1919, p. 149). The following seemingly convoluted paragraph on seed germination indicates the already advanced state of his evolutionary thought (Butler 1935):    

"The rose-seed did what it now does in the persons of its ancestors - to whom it has been so linked as to be able to remember what those ancestors did when they were placed as the rose-seed now is. Each stage of development brings back the recollection of the course taken in the preceding stage, and the development has been so often repeated, that all doubt - and with all doubt, all consciousness of action - is suspended. -- The action which each generation takes - [is] an action which repeats all the phenomena that we commonly associate with memory - which is explicable on the supposition that it has been guided by memory - and which has neither been explained, nor seems ever likely to be explained on any other theory than the supposition that there is an abiding memory between successive generations."


4. Heredity Equals Memory

Darwin's ideas were based on three fundamental principles - variation, heredity and natural selection (Darwin 1859). Variation caused an individual to differ in certain characters from others. Heredity caused characters to be transmitted from parents to children. Natural selection caused some individuals to survive and reproduce better than others. Of these three words - variation, heredity and natural selection - only the basis of the latter was understood. For the Victorians, heredity and variation were unknowns to be entrusted to researchers of future generations. The words "heredity" and "variation" were handles with which to manipulate these principles, irrespective of their underlying mechanisms.

    Initially heredity was seen as the process by which characters (which we would now call the phenotype) were passed forward from parent to child. However, unknown to Butler, in 1870 this perspective had been shifted by the physiologist Ewald Hering in Prague. Like Butler, Hering put the onus on the embryo, which remembers its parents. The newly formed embryo is a passive recipient of parental information (which we now call the genotype), and this information is used (recalled) by the embryo to construct itself (i.e. to regenerate phenotype from genotype). Heredity and memory are, in principle, the same. Heredity is the transfer of stored information. This powerful conceptual leap led to new territory. Evolutionary processes could thenceforth be thought of in the same way as mental processes. In a lecture delivered at the anniversary meeting of the Imperial Academy of Sciences in Vienna, Hering considered (Butler 1880, p. 131) that:

"We must ascribe both to the brain and body of the new-born infant a far-reaching power of remembering or reproducing things which have already come to their development thousands of times in the persons of their ancestors."

More simply, Butler (1985) wrote:

"There is the reproduction of an idea which has been produced once already, and there is the reproduction of a living form which has been produced once already. The first reproduction is certainly an effort of memory. It should not therefore surprise us if the second reproduction should turn out to be an effort of memory also."

    Hering wanted his strictly materialist position be understood (Butler 1880, p. 99): "Both man and the lower animals are to the physiologist neither more nor less than the matter of which they consist." He held that ideas that disappeared from consciousness would be stored in some material form in the unconscious memory, from which they could be recalled to consciousness. Thus, "the physiology of the unconscious is no 'philosophy of the unconscious'" (Butler 1880, p. 113).

    Hering considered that actions that were repeated were more likely to be stored in memory, and hence more accurately recalled, than actions that were performed only once or twice. So practice-makes-perfect with respect both to psychomotor functions, such as reproducing music on the piano, and to hereditary functions, such as reproducing a cell from a parent cell, and reproducing an organism from either a parent organism (asexual reproduction) or parent organisms (sexual reproduction) (Butler 1880, p. 124):

"But if the substance of the germ can reproduce characteristics acquired by the parent during its single life, how much more will it not be able to reproduce those that were congenital to the parent, and which have happened through countless generations to the organized matter of which the germ of today is a fragment? We cannot wonder that action already taken on innumerable past occasions by organized matter is more deeply impressed upon the recollection of the germ to which it gives rise than action taken once only during a single lifetime."

Among the recollections of the germ would be instinctual (innate) actions. As far as we know, a pianist, having practiced a piece so well that it can be played virtually unconsciously, does not pass knowledge of that piece through the germ line to children. Yet, Hering held that some psychomotor activities of comparable complexity are passed on (Butler 1880, p. 126):

"Not only is there reproduction of form, outward and inner conformation of body, organs, and cells, but the habitual actions of the parent are also reproduced. The chicken on emerging from the eggshell runs off as its mother ran off before it; yet what an extraordinary complication of emotions and sensations is necessary in order to preserve equilibrium in running. Surely the supposition of an inborn capacity for the reproduction of these intricate actions can alone explain the facts. As habitual practice becomes a second nature to the individual during his single lifetime, so the often-repeated action of each generation becomes a second nature to the race."

      Hering's lecture was noted with approval by the German evolutionist Ernst Haeckel (1876) in a book which was reviewed in Nature (Lankester 1876). Haeckel sent a copy to Charles Darwin, who forwarded it to his young research associate, the physiologist George Romanes (1848-1894; Darwin 1876). Butler learned of Hering's lecture in 1877 from Darwin's son, Francis (Jones 1919, p. 257), when telling him about the impending publication of Life and Habit, his first major evolution book (Butler 1878). This was to be warmly reviewed by Alfred Wallace in Nature (1879) as a "remarkable work," notable for its "originality" and "logical completeness," that was "in great part complementary" to Darwinian ideas. Unlike Hering, a scientist who had made a one-time sortie into the professionally hazardous area of evolutionary speculation, for the next two decades the polymathic Butler, while claiming disingenuously to "know nothing of science" (Butler 1878, p. 2), continued to develop his theme, frequently employing the technique of subject-object inversion of which he was a master. For example, Life and Habit contained the memorable phrase: "A hen is only an egg's way of making another egg" (Butler 1878, p. 134). In the twentieth century, zoologist Richard Dawkins was to argue similarly that a body is only a gene's way of making another gene (Dawkins 1976).

Butler's second major work, Evolution Old and New (1879), constituted a serious indictment of the failure of the Darwinists to acknowledge their intellectual antecedents - Comte Georges Louis Leclerc de Buffon (1707-1788), Doctor Erasmus Darwin (1731-1802), and Chevalier Jean Baptiste Pierre de Lamarck (1744-1829) - later summarized as: "Buffon planted, Erasmus Darwin and Lamarck watered, but it was Mr. Darwin who said, 'That fruit is ripe,' and shook it into his lap" (Butler 1887, p. 291). This, to put it mildly, did not please the Darwinians, particularly Romanes who, by virtue of his deep interest in evolution and mental function (Romanes 1884), was well prepared to understand Butler's work (Butler 1884). Charles Darwin, perhaps still smarting decades after being publicly accused of pococurantism by Butler's distinguished grandfather, was definitely not amused (Jones 1911).

5. Memory as Stored Information

Like other Victorians, Butler did not often use the word "information" in the context of heredity, but his third major work, Unconscious Memory (1880, p. 252), made clear his understanding of memory as stored information:      

"Does the offspring act as if it remembered? The answer to this question is not only that it does so act, but that it is not possible to account for either its development or its early instinctive actions upon any other hypothesis that that of its remembering, and remembering exceedingly well. The only alternative is to declare -- that a living being may display a vast and varied information concerning all manner of details, and be able to perform most intricate operations, independently of experience and practice." [My italics]

In his fourth major work on evolution, Luck or Cunning, Butler (1887, p. 308) outlined the concept of information and its symbolic representation in a form conditioned by factors both internal and external:

"As this idea [of a thing] is not like the thing itself, neither is it like the motions in our brain on which it is attendant. It is no more like these than, say, a stone is like the individual characters, written or spoken, that form the word 'stone' ... .The shifting nature ... of our ideas and conceptions is enough to show that they must be symbolic and conditioned by changes going on within ourselves as much as those outside us."

    In a lecture in 1894 on "Thought and Language" attacking the views of Oxford philologist Friedrich Max Muller (1823-1900), Butler was more poetic (Butler 1914b, 162):

"Some ideas crawl, some run, some fly; and in this case words are the wings they fly with, but they are only wings of thought or of ideas, they are not the thought or ideas themselves."

    While never using the word "information," in his "provisional theory of pangenesis" Darwin (1868) had written of "formative matter" and "formative elements," noting that "the child, strictly speaking, does not grow into the man, but includes germs which slowly and successively become developed and form the man." He equated "germs" with the "formative matter," which "consists of minute particles or gemmules." These were fundamental units each associated with a particular inherited character, and capable of independent multiplication by "self-division." Darwin's cousin, Francis Galton, when further developing this, touched on the idea of information and anticipated Weismann in postulating separation of germ-line and soma (Galton 1872; Galton 1876). Today we can best equate gemmules with genes (Forsdyke 2001, p. 151). Butler (1878, p. 187) supposed "a memory to 'run' each gemmule," but, of course, he knew the material basis of the storage of inherited information no more than he knew the material basis of the storage of mental information. We now know that DNA is the former, but still have little understanding of the latter.

    If challenged, Butler would probably have speculated that both forms of storage were the same (Occam's principle). Current evidence does not support this. Just as segments of DNA (genes) contain the information for the assembly, by some chain of events, of the components of, say, a hand, so segments of DNA should contain the information for a particular pattern of cerebral "wiring" (an "anatomical plan of the ganglia;" see below) that would, by some chain of events, be manifest as a particular instinct. Can such germ-line memories relate to memories acquired during individual lifetimes ("cerebral alterations")? A year before Darwin's death in 1882, George Romanes pointed out (Butler 1884, pp. 247-248):

"We can understand, in some measure, how an alteration in brain structure when once made should be permanent, -- but we cannot understand how this alteration is transmitted to progeny through structures so unlike the brain as are the products of the generative glands [gametes]. And we merely stultify ourselves if we suppose that the problem is brought any nearer to solution by asserting that a future individual while still in the germ has already participated, say, in the cerebral alterations of its parents."

     Butler seems to have been the first to extrapolate, from the principle that "higher" forms of life have evolved from "lower" forms, that eventually, beyond man, machines made by man would take over the planet (Butler 1914a, pp. 179-185). Today we would equate this with computers. Man may be only a computer's way of making another computer. Butler's genius was in seeing that inheritance and mental function, like today's computers, are all "modes of memory" (Butler 1878, pp. 299-300):      

"Life is that property of matter whereby it can remember. Matter which can remember is living; matter which cannot remember is dead. Life, then, is memory. The life of a creature is the memory of a creature. We are all the same stuff to start with, but we remember different things, and if we did not remember different things we should be absolutely like each other. As for the stuff itself, we know nothing save only that it is 'such as dreams are made of'." [Butler's italics]

In his Mental Evolution in Animals, Romanes (1884) was happy to consider instinctual activities, such as bird migration and nest-making, as requiring inherited ("ready-formed") information:

"Many animals come into the world with their powers of perception already largely developed. This is shown -- by all the host of instincts displayed by newly-born or newly-hatched animals. --The wealth of ready-formed information, and therefore of ready-made powers of perception, with which many newly-born or newly-hatched animals are provided, is so great and so precise, that it scarcely requires to be supplemented by the subsequent experience of the individual."[My italics]

     But, much to Butler's frustration, he would not move beyond this (Butler 1884, p. 236):

"Mr. Romanes -- speaks of 'heredity as playing an important part in forming memory of ancestral experiences;' so that whereas I want him to say that the phenomena of heredity are due to memory, he will have it that the memory is due to heredity --. Over and over again Mr. Romanes insists that it is heredity which does this or that. Thus, it is 'heredity with natural selection which adapt the anatomical plan of the ganglia;' but he nowhere tells us what heredity is any more than Messrs. Herbert Spencer, Darwin and Lewes have done. This, however, is exactly what Professor Hering, whom I have unwittingly followed, does. He resolves all phenomena of heredity, whether in respect of body or mind, into phenomena of memory. He says in effect, 'A man grows his body as he does, and a bird makes her nest as she does, because both man and bird remember having grown body and made nest as they now do -- on innumerable past occasions. He thus reduces life from an equation of say 100 unknown quantities to one of 99 only, by showing that heredity and memory, two of the original 100 unknown quantities, are in reality part of one and the same thing." [Butler's italics]


6. Variation

"Memory" refers either to the physiological process by which stored information is retrieved, or to the actual stored information itself. In the former sense, the storage of information is implicit. In the latter sense, memory is stored information. Whatever the sense, the concepts of memory and of stored information are very close. At some point in time information is selected and assigned to a store, and at a later point in time it may be retrieved. But storage of information is not enough. Information must be safely stored. To select and to store is not to preserve. Stored information must also be preserved information. When preservation is not perfect, there is variation. When there is variation, there can be both linear and branching evolution. Thus, in 1966 the US biologist George C. Williams defined a gene, not as a unit of function (observed as some phenotypic characteristic), but as a unit of preservation. This, and the related work of the Japanese biophysicist Akiyoshi Wada, are discussed elsewhere in much technical detail (Forsdyke 2004a, b; Lee, Mortimer and Forsdyke 2004; Forsdyke 2006).

      Apparently unaware of Joseph Hooker's "doctrine of creation by variation" (Hooker 1860, p. 315), Butler (1878, p. 263) declared that "the 'Origin of Variation,' whatever it is, is the only true 'Origin of Species'". However, he knew the underlying mechanisms by which Nature's expressions vary from generation to generation, no more than he knew the underlying mechanisms by which mental expressions vary from day to day. Nor did Butler know the scope of variation. Variation can only occur in that which already exists, and the nature of that which already exists, limits the scope of its variation. Just as the scope of a dice is limited by its structure, so the scope of variation of a living form is limited. From time to time a human is born with six fingers, but, as far as we know, no human has ever been born (or even conceived) with a hundred fingers, or with feathers (Bateson 1894).

      Butler often wrote of dice (e.g. see below). The variation in landing position of a balanced dice is random. The variation in landing position of an unbalanced dice is non-random. So, from first principles, it might be supposed that variation in living forms would be either a chance event, or biased. Whether unbiased or biased, a variation might be beneficial, deleterious, or somewhere in between. Within its scope, the dice of life might be multifaceted allowing fine gradations in variation that might appear continuous. Alternatively variations might be quite discrete (e.g. the usual six-sided dice), so appearing discontinuous. And would the dice of life be thrown? If so, by what agency? If not thrown, then any variation that occurred would be "spontaneous" and perhaps unbiased. If the dice of life were thrown, then there would be a thrower. Would this agency be internal or external to the organism? In either case, could the agency direct the bias to the adaptive advantage of the organism? In other words, could the agency "design" the organism?

       Among these alternatives, current evidence shows that variation is a spontaneous property of matter, but that it may be affected, both quantitatively and qualitatively, by external agents, such as radiation (Forsdyke 2001; 2006). Within its scope, variation is unbiased.  It is not directed. There is no design by means of variation, either internal or external. If an organism is closely adapted to its environment then a variation is unlikely to be beneficial. If the environment has recently changed dramatically, then a variation may more likely be beneficial. But variations, per se, do not occur "for the good of the organism" (teleology).

      However, the parallels he had drawn between memory and heredity suggested to Butler a set of alternatives that had some plausibility at the time. He opted for an agency internal to the organism that would, in small steps, bring about variations that would accumulate to the advantage of the organism. This set of alternatives had gained some scientific respectability both in England (e.g. Spencer), and in continental Europe (e.g. Karl Wilhelm von Nageli) and some came to call it "orthogenesis," implying an innate tendency to progressive development (Kellogg 1908). Butler's teleology rested on grounds that were both aesthetic and logical.

's argument that natural selection would cruelly send the weaker to the wall and select the fittest to survive sufficed "to arouse instinctive loathing; -- such a nightmare of waste and death is as baseless as it is repulsive" (Butler 1914b, p. 293). To buttress this feeling, Butler argued along the lines of Ernst Brucke (1861) who had viewed unicellular organisms as elementary versions of organisms considered higher in the evolutionary scale. Microscopic studies of unicellular organisms, such as amoebae, showed them to possess organelles analogous to the organs of multicellular organisms. Thus, an amoeba, far from being an amorphous mass of protoplasm, was seen to extrude "arms" (pseudopodia), fashion a "mouth", and digest it prey in a prototypic stomach (digestive vacuole). If it could achieve this degree of sophistication, then perhaps it could, in an elementary way, also think? Butler (1879, p. 40) was quite open with his premises:

"Given a small speck of jelly with some power of slightly varying its actions in accordance with slightly varying circumstances and desires - given such a jelly speck with a power of assimilating other matter, and thus, of reproducing itself, given also that it should be possessed of a memory and a reproductive system --".[my italics]

When it served his purpose Butler (1914b, p. 301) often appealed to "people in ordinary life," or to "plain people":

"The difference between Professor Weismann and, we will say, Heringians consists in the fact that the first maintains the new germ-plasm, when on the point of repeating its developmental process, to take practically no cognisance of anything that has happened to it since the last occasion on which it developed itself; while the latter maintain that offspring takes much the same kind of account of what has happened to it in the persons of its parents since the last occasion on which it developed itself, as people in ordinary life take things that happen to them. In daily life people let fairly normal circumstances come and go without much heed as matters of course. If they have been lucky they make a note of it and try to repeat their success. If they have been unfortunate but have recovered rapidly they soon forget it; if they have suffered long and deeply they grizzle over it and are scared and scarred by it for a long time. The question is one of cognisance or non-cognisance on the part of the new germs, of the more profound impressions made on them while they were one with their parents, between the occasion of their last preceding development and the new course on which they are about to enter."

      Thus, Butler downplayed chance ("luck") and championed an intrinsic capacity for bias ("cunning") as the means by which advantageous characters acquired by parents would be transmitted to their children. In short, he appealed to the doctrine of the inheritance of acquired characters that had been advocated in France by Lamarck (1809). Butler even went so far as to assert, in Romanes' words (Butler 1884, p. 248), "that a future individual while still in the germ has already participated -- in the cerebral alterations of its parents." However, like Hering, Butler maintained a strictly materialist position. He used the words "intelligent" and "design," often separately, and sometimes together (Butler 1879, p. 11), but never in a way as to suggest the involvement of an agency external to the organism (Butler 1914b, p. 253-254):

"The two facts, evolution and design, are equally patent to plain people. There is no escaping from either. According to Messrs. Darwin and Wallace, we may have evolution, but are in no account to have it mainly due to intelligent effort, guided by ever higher and higher range of sensations, perceptions and ideas. We are to set it down to the shuffling of cards, or the throwing of dice without the play, and this will never stand. According to the older men [e.g. Lamarck], the cards did count for much, but play was much more. They denied the teleology of the time - that is to say, the teleology that saw all adaptation to surroundings as part of a plan devised long ages since by a quasi-anthropomorphic being who schemed everything out much as a man would do, but on an infinitely vaster scale. This conception they found repugnant alike to intelligence and conscience, but, though they do not seem to have perceived it, they left the door open for a design more true and more demonstrable than that which they excluded."

      Butler soared beyond the comprehension of most of his contemporaries on the wings of his conceptual insight that heredity was the transfer of stored information. The German zoologist Richard Semon (1859-1918) acknowledged Butler's influence in developing his "mneme" theory of memory (Hartog 1914; Russell 1916; Richards 1987; Schachter 2001). Dawkins (2003) was unaware of this when he suggested the word "meme" as a self-replicating element of culture, passed on by imitation. Although Butler's dalliance with Lamarckism has not won support, we recognize today that the robustness of the underlying idea led him closer to solutions to fundamental biological problems such as the origin of sex, the sterility of hybrids, and aging (Butler 1878, p. 296).

      For example, since parental gametes can transfer information to offspring only while the parents are reproductively active (i.e. young), Butler deduced that offspring would receive no information from parental gametes to help them cope with aging. Today, we consider this to be true, but for entirely different reasons - namely, that it is natural selection acting on the young, not on the old, that increases genetic fitness (as measured by number of descendents produced). Over evolutionary time, natural environmental hazards have often eliminated organisms before their reproductive lives are over. Thus, the germ-line normally does not contain information for extending lifespan long after the mean age of reproduction. Exceptions may occur in the case of species where grandparental care of offspring is important (e.g. man) and in species that, for various adaptive reasons have incidentally developed additional ways of protecting DNA against damages that can accelerate aging (e.g. a tortoise shell protects against predators, but should also be a very effective shield against natural radiation; Bernstein and Bernstein 1991).          

     For his time, Butler's correct equation of heredity with memory (i.e. the transfer of stored information) was beyond the pale, but his incorrect Lamarckism seemed plausible. Herbert Spencer was a strong supporter of this viewpoint and, at Darwin's behest, Romanes spent several years fruitlessly seeking to show that gemmules containing information for acquired characters could be transferred from normal parental tissues (soma) to the gonads (germ-line; Romanes 1896; Biggers 1991). And as late as 1909 Haeckel was proclaiming that "natural selection does not of itself give the solution of all our evolutionary problems. It has to be taken in conjunction with the transformism of Lamarck, with which it is in complete harmony".


7. Lamarckism       

In Scotland a professor of engineering, Fleeming Jenkin, noted that for "an impartial looker-on" the best way to question "the Darwinians" was to "admit the facts, and examine the reasoning" (Jenkin 1867). This seems to have been the approach taken by Butler. He did not himself read the primary literature. He relied on the facts provided by the Darwinians and their few opponents, especially Mivart (1871) (Butler 1887, p. 5). Although Butler was greatly disillusioned on finding that the men of science had not done their home-work regarding early sources (Butler 1879), he did not extrapolate this to the current literature. He assumed that they were conversant with the current literature and, where deemed relevant, made it known in their publications. He identified "one apparently very important problem which I do not at this moment see how to connect with memory, namely, the tendency on the part of offspring to revert to an earlier impregnation" (e.g. to appear more like the grandparents than the parents; Butler 1878, pp. 182-183). However, it is likely he did not know of the private correspondence between Darwin, Thomas Huxley and Joseph Hooker where they discussed Charles Naudin's studies of plant hybrids in France (Naudin 1856; Forsdyke 2001. p. 24). These indicated that parental characters might be inherited discretely (discontinuously), rather than blending (continuously) in their offspring. Thus, after a period of latency the characters might emerge unchanged in future generations, even though appearing to be absent in immediate offspring. Probably privy to this correspondence, Romanes later wrote to a friend (1894):

"I have found after several years experimenting with rabbits, rats and [undecipherable], that one may breed scores and hundreds of first crosses between different varieties, and never get a single mongrel throwing intermediate characters - or indeed any resemblance to one side of the house. Yet, if the younger are subsequently crossed inter se (i.e. brothers and sisters, or first crossings) the crosses parentage at once repeats itself. Ergo, even if the pups wh[ich] are to be born appear to give a negative result, keep them to breed from with one another."

The prevailing view was that parental characters were blended in their offspring. Jenkin (1867), for example, saw members of a biological species as enclosed within a sphere and noted that there would always be a tendency for individuals to vary centripetally, rather than centrifugally:

"A given animal -- appears to be contained -- within a sphere of variation; one individual lies near one portion of the surface; another individual, of the same species, near another part of the surface; the average animal at the centre. Any individual may produce descendents varying in any direction, but is more likely to produce descendents varying towards the centre of the sphere, and the variations in that direction will be greater in amount than the variations towards the surface. Thus a set of racers of equal merit indiscriminately breeding will produce more colts and foals of inferior than of superior speed, and the falling off of the degenerate will be greater than the improvement of the select."

Given blending inheritance, it was difficult to see how a rare new variant organism, even if personally advantaged by virtue of the variation, would find a mate with a similar variation. Rather, the variant (e.g. white) would be swamped by crossing with the abundant non-variant type (e.g. black) to produce a blend (e.g. grey). With further crossing with the non-variant type (black) the descendents of the variant would soon be indistinguishable from the original non-variant type. This latter point was emphasized by Jenkin (1867):

"Any favourable deviation must -- give its fortunate possessor a better chance of life; but this conclusion differs widely from the supposed consequence that a whole species may or will gradually acquire some one new quality, or wholly change in one direction and in the same manner."

    Easy access to a mate with the same mutation had been a strength of the Lamarckist doctrine of the inheritance of acquired characters. According to Lamarck (and Butler) organisms subjected to the same environmental provocation would simultaneously adapt and so be in possession of the same variant character. Being colocalized in the same environment there would then be little difficulty in their finding a mate with the same adaptation (Butler 1879, pp. 342-343, 1914b, p. 250). But, unknown to most of the men of science (and hence to Butler), by 1865 Mendel had obtained good evidence for discontinuous, non-blending, inheritance.  Knowledge of Mendel's breeding studies with peas, and of Michael Guyer's studies of meiotic chromosomes, emerged in 1900 only two years before Butler's death (Bateson 1909; Bungener and Buscaglia 2003). Thus, although Butler was probably never aware of it, his arguments rested on unstable Lamarckist foundations.

There was also an anti-Lamarckian argument that Butler had acknowledged, but never satisfactorily dealt with (Butler 1878, p. 237). Among colonial insects there is often a much greater separation of the germ line from the soma than in other organisms. For example, the germ line is personalized in the form of the queen bee. The soma is personalized in the forms of the neuter worker bees. Through "use or disuse" the latter might come to strengthen or deplete characters in a Lamarckian fashion. But how could the corresponding Darwinian "gemmules" transfer back to their queen. Apart from recent work showing the systemic spread of RNA within a plant, which is probably an antiviral defense (Waterhouse, Wang and Lough 2001; Liu 2006), there is no evidence that gemmules (i.e. genes) can transfer from soma to germ line within a body, let alone between bodies (that do not interact sexually). Many of the phenomena associated with colonial insects have found satisfactory explanations in terms of modern selfish-gene theory (Dawkins 1976; 2003). The possibility of transgenerational inheritance involving transfer of epigenetic information is considered elsewhere (Forsdyke 2006, p. 323).


8. Recent Controversy over "Intelligent Design"

In the wake of various lawsuits over the teaching of Darwinian evolution, a major proponent of the so-called "intelligent design" (ID) alternative was recently given the opportunity to clarifying the issue in the "op-ed" section of The New York Times. It seemed possible that, once and for all, the "widespread confusion about what intelligent design is and what it is not" would be dispelled, and the case that ID "is not a religiously based idea" would be set out with irrefutable clarity. Sadly, the old arguments of the eighteenth century theologian William Paley, a distant relative of Butler, were dutifully repeated. Organic forms were just too complex to have arisen without some form of cognitive input, which to Paley, but apparently not to the ID movement, meant "God" (Behe 1996; Behe 2005). Not surprisingly, a critical response was provoked from the President of the US National Academy of Sciences (Alberts 2005):

"In evolution, as in all areas of science, our knowledge is incomplete. But the entire success of the scientific enterprise has depended on an insistence that these gaps be filled by natural explanations, logically derived from confirmable evidence. Because 'intelligent design' theories are based on supernatural explanations, they can have nothing to do with science."

Simply put, one party, having defined its own version of ID, declared that it was not religious, and so was scientific. The other party declared that all ID theories were religious, and so were not scientific. Neither party appeared aware that in the nineteenth century Samuel Butler had furnished Paley's argument with a non-supernatural, and what then appeared, a scientifically coherent, foundation. Since the balance of evidence turned strongly against it, Butlerian ID did not gain support. In particular, there was the doctrine of the non-inheritance of acquired characters, with separation of the cells of the germ line from those of the mortal soma, as argued by August Weismann (1904). However, Butlerian ID was a respectable scientific hypothesis in accord with the canons of its time. Butler's evolutionary ideas won the praise of Francis Darwin (1908; see also Jones 1911), the Cambridge biologist William Bateson (1909, p. 88), and other scientists (Hartog 1914; Russell 1916).


9. Conclusions

Butlerian intelligent design must be clearly distinguished from the "intelligent design" that is currently a subject of much controversy. Butlerian intelligent design had a coherent foundation that was not "based on supernatural explanations," and is part of the history of science. It can, and should, form part of science history curricula. Modern proponents of "intelligent design" do not invoke Butlerian roots, and provide no coherent alternative. The concept of information in the context of hereditary transmission has ancient roots (Delbruck 1971), and in 1863 the geologist Charles Lyell devoted an entire chapter to the subject. Darwin's (1868) pangenesis touches upon it. But the formulations of Hering and Butler go far beyond this. Despite fundamental errors, their nineteenth century views have an intrinsic robustness and may prove illuminating as in the twenty-first century the biosciences become increasingly information-based (Baldi 2001; Barbieri 2003; Yockey 2005; Forsdyke 2006).   


I thank Claus Emmeche for a helpful review. Queen's University hosts my web-pages where further background and full-text versions of some of the references may be found. Macmillan Reference Ltd. kindly placed my brief biographies of Mendel and Romanes online in the Nature Encyclopedia of Life Sciences, now made available by Wiley ( The Project Gutenberg has made many of Butler's works available online (


Alberts, Bruce

2005: "Intelligent Design", The New York Times, February 12.

Baldi, Pierre

2001: The Shattered Self: The End of Natural Evolution, Cambridge, MA: MIT Press.

Barbieri, Marcello

 2003: Organic Codes, Cambridge: Cambridge University Press.

Bateson, William

   1894: Materials for the Study of Variation Treated with Especial Regard for   Discontinuity in the Origin of Species, London: Macmillan.

      1909: "Heredity and Variation in Modern Lights", in A C Seward (ed), Darwin and Modern Science, Cambridge: Cambridge University Press, pp. 85-101.

Behe, Michael J.

       1996: Darwin 's Black Box. The Biochemical Challenge to Evolution, New York: The Free Press.

      2005: "Design for Living", The New York Times, February 7.

Bernstein, Carol, and Harris Bernstein

      1991: Aging, Sex and DNA Repair, San Diego: Academic Press, pp. 4-11.

Biggers, J. D.

      1991: "Walter Heape, FRS: a pioneer in reproductive biology. Centenary of his embryo transfer experiments. Journal of Reproduction and Fertility 93, 173-186.

Bowlby, John

      1990: Charles Darwin. A New Life, New York:Norton, p. 334.

Brucke, Ernst

      1861: "Die Elementarorganismen", Sitzungsberichte der Akademie der Wissenschaften Wein, Mathematische-wissenschaftliche Classe 44, 381-406.

Bungener, Patrick, and Marino Buscaglia

      2003: "Early connection between cytology and Mendelism: Michael F. Guyer's contribution. History and Philosophy of the Life Sciences 25, 27-50. [See also Guyer in these web-pages (Click Here)]

Butler, Samuel

      1878: Life and Habit, London:Trubner.

      1879: Evolution, Old and New, London:Hardwicke and Bogue.

      1880: Unconscious Memory, London:Bogue.

      1884: Selections from Previous Works, with Remarks on Mr. G. J. Romanes' "Mental  Evolution in Animals," and a Psalm of Montreal , London:Trubner, pp. 228-254.

      1887: Luck or Cunning as the Main Means of Organic Modification? London:Trubner.

      1914a: A First Year in Canterbury Settlement with Other Earlier Essays, R. A. Streatfeild (ed), London: Fifield, pp. 149-185.

      1914b: The Humour of Homer and Other Essays, New York:Kennerley, pp. 209-313.

      1935: Erewhon or Over the Range, Harmondsworth: Penguin Books, pp. 197-198.

      1985: The Notebooks of Samuel Butler, H. Festing Jones (ed), London: Hogarth Press, p. 59.

Darwin, Charles

      1859: The Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life, London:John Murray.

      1867: Letter to Canon Farrar, March 5, in Francis Darwin (ed), More Letters of  Charles Darwin, Volume 2, New York : Appleton (1903), pp. 441-442.  

      1868: The Variation of Animals and Plants under Domestication, London: John Murray, pp. 369-399.

      1876: Letter to Romanes, May 29, in Francis Darwin (ed), More Letters of Charles Darwin, Volume 1, New York: Appleton (1903), pp. 363-365.

Darwin, Francis

      1908: Address to the British Association for the Advancement of Science, Dublin, London: The British Association, pp. 14, 22.

Dawkins, Richard

      1976: The Selfish Gene, Oxford: Oxford University Press.

      2003: A Devil's Chaplain, Boston: Houghton Mifflin, p. 252.

Delbruck, Max

      1971: "Aristotle-totle-totle", in Jaques Monod and Felix Borek (eds), Of Microbes and Life, New York: Columbia University Press, pp. 50-55.

Forsdyke, Donald R.

      2001: The Origin of Species, Revisited, Montreal: McGill-Queen's University Press.

      2004a: "Chromosomal speciation: a reply", Journal of Theoretical Biology 230, 189-196.

      2004b: "Regions of relative GC% uniformity are recombinational isolators", Journal of Biological Systems 12, 261-271.

      2006: Evolutionary Bioinformatics, New York: Springer.

Galton, Francis

      1872: "On blood relationship," Proceedings of the Royal Society 20, 394-402.

      1876: "A theory of heredity", Journal of the Anthropological Institute 5, 329-348.

Haeckel, Ernst

      1876: Die Perigenesis der Plastidule oder die Wellenzeugung der Lebenstheilchen, Berlin:Reimer, pp. 40-4, 67, 77.

      1909: "Charles Darwin as an Anthropologist", in A. C. Seward (ed), Darwin and Modern Science, Cambridge: Cambridge University Press, pp. 137-151.

Hartog, Marcus

      1914: "Samuel Butler and recent mnemic biological theories", Scientia 15, 38-52.

Hering, Ewald

      1870: Uber das Gedachtniss als eine allgemeine Function der organisirten Materie. Vienna: Karl Gerold's Sohn. ("On memory as a universal function of organized matter." A lecture delivered on May 30th at the anniversary meeting of the Imperial Academy of Sciences at Vienna, translated by Butler "with the kind assistance of friends" in his 1880: Unconscious Memory, London: David Bogue, pp. 97-133. (Another, anonymous, translation, with some interesting differences, is in the 1896 Religion of Science Library issue cited next).

      1896: On Memory and the Specific Energies of the Nervous System, Chicago: The Open Court Publishing Company for The Religion of Science Library.

Hooker, Joseph Dalton

      1860: "On the origination and distribution of vegetable species: introductory essay to the flora of Tasmania", American Journal of Science and Arts 29, 305-326.

Iltis, Hugo

      1932: Life of Mendel, London:Allen and Unwin, p. 103.

Jenkin, H. C. Fleeming

      1867: "The origin of species", The North British Review 46, 277-318.

Jones, Henry Festing

      1911: Charles Darwin and Samuel Butler. A Step Towards Reconciliation. London: Fifield.

1919: Samuel Butler Author of Erewhon (1835-1902). A Memoir, Vols. 1 and 2. London: Macmillan.

Kay, Lily E.

      2000: Who Wrote the Book of Life? A History of the Genetic Code, Stanford: Stanford University Press.

Kellogg, Vernon L.

      1908: Darwinism Today. A Discussion of Present-Day Scientific Criticism of the Darwinian Selection Theories, Together with a Brief Account of the Principal other Proposed Auxiliary and Alternative Theories of Species-Forming, New York:Holt, pp. 274-289.

Lamarck, Jean-Baptiste

      1809: Philosophie Zoologique. Translated by Hugh Elliot as Zoological Philosophy, Chicago:University of Chicago Press, 1984.

Lankester, Edwin Ray

      1876: "Perigenesis versus pangenesis - Haeckel's new theory of heredity", Nature 14, 235-238.

Lee, Shang-Jung, James R. Mortimer, and Donald R. Forsdyke

      2004: "Genomic conflict settled in favour of the species rather than the gene at extreme GC percentage values", Applied Bioinformatics 3, 219-228.

Liu, Yongsheng

2006: "Historical and modern genetics of plant graft hybridization", Advances in Genetics 56, 101-129.

Lyell, Charles

      1863: The Geological Evidences of the Antiquity of Man with Remarks on Theories of the Origin of Species by Variation. Philadelphia: G. W. Childs, pp. 454-470.

Mendel, Gregor Johann

      1865: "Versuche uber Pflanzen Hybriden", Verhandlung des naturforschenden Vereines in Brunn 4, 3-47.

Mivart, St. George J.

      1871: On the Genesis of Species, London:Macmillan.

Naudin, Charles

     1856: "Observations constatant le retour simultane de la descendance d'une plante hybride aux types paternels et 
      maternels", Compte Rendu Academie Sciences, Paris 42, 628-636.

Orel , Vítĕzslav

      1984: Mendel, Translated by S. Finn, Oxford: Oxford University Press, p. 69.

Richards, Robert J.

      1987: Darwin and the Emergence of Evolutionary Ideas of Mind and Behaviour. Chicago: Chicago University Press.

Roberts, H. F.

1929: Plant Hybridization before Mendel. Princeton: Princeton University Press.

Romanes, Ethel

      1896: The Life and Letters of George John Romanes. London: Longmans, Green and Co., pp. 19-50, 223.

Romanes, George John

      1884: Mental Evolution in Animals, with a Posthumous Essay on Instinct by Charles Darwin, New York: Appleton , p. 131.

1894: Letter to Schafer, May 18, London: Welcome Museum of the History of Medicine.[Romanes died a few days later]

Russell, Edward Stuart

      1916: Form and Function, London: Murray, pp. 335-344.

Schachter, Daniel L.

      2001: Forgotten Ideas, Neglected Pioneers: Richard Semon and the Story of Memory, Psychology Press, Philadelphia.

Smith, John Maynard

      1999: "The idea of information in biology", The Quarterly Review of Biology 74, 395-400.

2000: "The concept of information in biology", Philosophy of Science 67, 177-194.

Wallace, Alfred Russel

      1879: "Organization and intelligence", Nature 19, 477-480.

Waterhouse, Peter M., Ming-Bo Wang, and Tony Lough

      2001: Gene silencing as an adaptive defence against viruses. Nature 411, 834-841.

Weismann, August

      1904: The Evolution Theory, Volume 1, London: Edward Arnold, p. 411.

Williams, George C.

      1966: Adaptation and Natural Selection, Princeton: Princeton University Press.

Yockey, Hubert P.

       2005: Information Theory, Evolution, and the Origin of Life, Cambridge: Camb. Univ. Press. 

End Note on Dreams (Jan 2007)

1. Prospero in The Tempest declares "We are such stuff as dreams are made on."

2. In Erewhon Revisited (1901), Butler writes: "I wish some one would write a book about dreams and parthenogenesis -- for that the two are part and parcel of the same story -- a brood of folly without father bred -- I cannot doubt." In his notebooks (Further Extracts from the Notebooks of Samuel Butler 1934), Butler declared: "Dreams are a mode of parthenogenesis when the vibrations pre-existing in the brain vivify up to a certain point but do so all wrong as though in a mental ovarian tumour, through want of fecundation by incoming vibrations from exterior objects."

3. Gregory Bateson in Mind and Nature (1979) takes this as Butler arguing: "As dreams are to thought, so parthenogenesis is to sexual reproduction. Thought is stabilized and tested against the template of external reality, but dreams run loose. Similarly, parthenogenesis can be expected to run loose; whereas zygote formation is stabilized by the mutual comparison of gametes."

End Note on Pernicious Anaemia (Apr 2010)

Butler was born on 4th December 1835 and died of pernicious anaemia on 18th June 1902 at the age of 66. 18 years later George Hoyt Whipple (1878-1976; right) found that raw liver was curative. George Minot and William Murphy later showed the active principle was in liver juice, but it was not until 1948 that the active principle, Vitamin B12, was identified. In 1934 Whipple, Minot and Murphy received the Nobel Prize for Physiology or Medicine. An independent observer has pointed out that, in appearance, GHW is remarkably like the author of these webpages (in a more sober mood). 

George Hoyt Whipple (1878-1976)


End Note (July 2010) - Commentary by Shaw

George Romanes crossed swords with Samuel Butler who, way ahead of his time, viewed heredity as information transmission from generation to generation. George Bernard Shaw counted himself as among "the best brains" that supported Butler. Shaw outlived them all. In an introductory essay to a reprinting of Butler's The Way of All Flesh, (1936; Oxford University Press) Shaw concluded:

Darwin, a simple-souled naturalist with no comprehension of the abyss of moral horror that separated his little speciality of Natural Selection from Butler's comprehensive philosophic conception of Evolution, may be pardoned for his foolish estimate of Butler as 'a clever unscrupulous man,' and for countenancing the belittling of him by Huxley and Romanes that now seems so ridiculous. They really did not know any better.


End Note (Nov 2010) - Darwin's Notebooks

One searches in vain for the I-word -- Information -- among Charles Darwin's Notebook writings (circa 1840) that were inspired by the early 19th century literature he was deeply reading. Yet those he read, such as Edmund Blyth (1837. Magazine of Natural History), sometimes used the I-word [Notebook C]: "Whereas the human race is compelled to derive the whole of its information through the medium of the senses, the brute is, on the contrary, supplied with an innate knowledge of whatever properties belong to all the natural objects around, which can in anywise affect its own interests or welfare." The M-word -- Memory -- being stored information, occurs often; e.g.[Notebook M]: "Now if memory << of a tune & words can thus lie dormant, during a whole life time, quite unconsciously of it, surely memory from one generation to another, also without consciousness, as instincts are, is not so very wonderful." Occasionally Darwin used the K-word - Knowledge -- as when he considered the power of limb regeneration possessed by some animals [Notebook D]:

I consider gemmation [budding] as artificial division. -- On this view each particle of animal must have structure of whole comprehended in itself. -- it must have the knowledge how to grow, & therefore to repair wounds -- but it has nothing to do with generation. --- Why crab can produce claw. but man not arm. hard to say -- if it were possible to support [keep alive] the arm of Man, when cut off, it would produce another man. -- That the embryo the thousandth of an inch should produce a Newton is often thought wonderful. it is part of the same class of facts that the skin grows over a wound.

Having read Charles Bell (1833) on insect development, Darwin commented [Notebook E]: "Seems to argue, that as the transformations from the egg, or larva, or foetus to perfect animal are adapted by foreknowledge, so must the mutations of species.!!"

    At that time Darwin pondered the relative roles of Lamarckian evolution and natural selection [Notebook N]: "An habitual action must some way affect the brain in a manner which can be transmitted. -- this is analogous to a blacksmith having children with strong arms. -- The other principle of those children. which chance? produced with strong arms, outliving the weaker ones, may be applicable to the formation of instincts, independently of habits. -- the limits of these two actions either on form or brain very hard to define." 

    From this we see that Butler was crossing ground that Darwin had already explored, yet Butler emerged with quite different conclusions. 

End Note (Jan 2013) - Haeckel's Support of Semon

As set out in detail in the paper on Butler and long-term memory (click below), the Butler-Hering viewpoint was adopted by Semon with little acknowledgement. Francis Darwin praised Semon while well-aware of Butler's work, but Haeckel was possibly unaware of Butler. At the Darwin celebration in 1909, Bateson praised Butler, while Haeckel referred to "This important Mneme-theory of Semon and the luminous physiological experiments and observations associated with it:"

The most important advance that evolution has made since Darwin and the most valuable amplification of his theory of selection is, in my opinion, the work of Richard Semon: Die Mneme als erhaltendes Prinzip im Wechsel des organischen Geschehens. He offers a psychological explanation of the facts of heredity by reducing them to a process of (unconscious) memory. The physiologist Ewald Hering had shown in 1870 that memory [stored information] must be regarded as a general function of organic matter, and that we are quite unable to explain the chief vital phenomena, especially those of reproduction and inheritance, unless we admit this unconscious memory.



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