Chapter 1

Darwinism on the Deathbed, 1870-1900: The Failings of Natural Selection

A thorough scientific history of biology in the first few decades after Darwin has yet to be written. Despite the plethora of studies in this field it remains the considered opinion of a recent authority that not one of them has succeeded in being other than 'desultory' and that all their labours have barely reduced the 'area demanding systematic exploration'.¹ Since the present study cannot pretend to offer even a rudimentary history of the biological sciences over the period in question, there is certainly no intention here of mobilising one view of the history of Darwinism* against another, or of discriminating amidst competing claims in any lengthy way. What is necessary, however, is that the particular bias of the following pages towards one reading of the progressive absorption of Darwinism into Victorian intellectual culture should be stated fairly exactly, since on this reading rests some part of the argument to be pursued. One might begin, then, with what can fairly be termed the traditional description of the fortunes of the Darwinian revolution. According to such accounts, which tend to be fond of the imagery of military strategy, the publication of The

Origin of Species began a campaign of reason versus irrationality in which a small platoon of Darwinians marched from strength to strength, quickly sweeping aside a feeble opposition of ageing High churchmen. It is traditional, too, for the latter to be vilified as the fit progeny of their illustrious forebear who declined to put his eye to Galileo's telescope. This kind of description has been offered times without number, but never more ably than by Julian Huxley.² The bare bones of the sequence he offers are as follows: that after 1859, over a span of some thirteen years, Victorian England took Darwin to its heart; from the early 1870s until close to the end of the century the romance was in full bloom; then an estrangement is said to have set in for a couple of decades with the rise of anti-Darwinian evolutionary theory, but by 1925 this had passed by. Neo-Darwinism of a rigour hardly entertained by Darwin himself has held sway ever since. A similar dating appeals to William Coleman, too, for he argues that by the turn of the century 'no serious alternative, other than the ever-dubious environmental (called neo-Lamarckian) hypotheses, could be seriously maintained'.³ Another historian of science, describing attitudes in this same period of the late 1890s, concurs with this: 'the climate of opinion was by no means wholly anti-Darwinian in character. Among the strongest supporters of natural selection were three of the most influential names in biology at the turn of the century: A.R. Wallace, John George Romanes [sic], and August Weismann.'⁴

It is worthwhile taking up this latter judgement for a moment, for in every detail it is, as a description of the unanimity of views of the inner Darwinian party, a misleading oversimplification which it is vital to clear up at once. Certainly it is unquestionable that Alfred Russel Wallace, the co-presenter of the natural selection hypothesis in 1858, adhered throughout his long career to the very strictest Darwinism as providing a sufficient explanation for the full variety of life on earth. Even as early as 1870, however, he was already insisting vehemently on one most notable exception to the all-sufficiency of Darwinism: Wallace excluded human evolution. Wallace's teleological* evolutionism, which he espoused (publicly at least) on the strictest empirical grounds, will be taken up on a later page. Here

it must suffice to point out that right from its inception Darwinism was prone to heresies from within its own order, as here with Wallace's reservation (stated sufficiently bluntly even in the late 1860s to give Darwin some pain) that natural selection is a power completely impotent to explain the prodigiously rapid evolution of our species. G.J. Romanes was no more of a Darwinjan in his later years. Though he admired Darwin personally to the point of adulation, he abandoned after Darwin's death in 1881 whatever faith he had had in the universality of Darwinism. In a survey, for example, five chapters long of the second volume of his Darwin, and after Darwin (1892-7) -- a survey which is surely the most extensive ever made of the evidence relating to teleological and dysteleological evolutionary mechanisms -- Romanes, very much closer to the situation than we are, states categorically that at the time of Darwin's death Wallace, even with his advocacy of natural selection restricted to the animal world, 'was absolutely alone in maintaining this opinion: the whole body of scientific thought throughout the world being against him'.⁵ Nor is Romanes' own position left in any doubt. At the end of his massive consideration, he ponderously allies himself with the majority with all the force of a man distilling decades of thought and experiment into a few phrases:

there are certain general considerations, and certain particular facts, which appear to render it probable that [various teleological factors] have played a highly important part in the process of organic evolution as a whole... there must be some other, though hitherto undiscovered, principle at work, which co-operates with natural selection.

And we find further that his anti-Darwinian sentiments formed, or began to form, much earlier. On 30 December 1883, in a private letter to Asa Gray, Romanes may be discovered thoughtfully toying with Wallace's by then firm conviction that the human brain must, on the anatomical evidence, have evolved from the simian under the direct guidance of 'a higher and supreme mind'.⁶ In the face of these considered and final remarks it is difficult to see how Romanes can properly be maintained in his place as one of the Darwinian triumvirate. As for the third member, August Weismann's uncompromising neo-Darwinism had only a few years in favour in the early 1890s and had certainly fallen out of the centre of disputation before the decade and the century were over. Starting off as a simple and luminous idea -- that the germ plasm, the physical principle of heredity, cannot be violated by the environment -- Weismannism quickly became ornamented with qualifications and swamped by internal inconsistencies as it tried to deal with the criticisms levelled at it. The formerly indivisible germ plasm was splintered into no less than nine fragments (molecules, biophores, determinants, ids, idants, idioplasm, somatic-idioplasm, morphoplasm and apical-plasm; some of them externally influenced) which Romanes said drily reminded him of the Circles of the Inferno, in that from those regions regions one returned to the firm footing of science with a sense of having indeed been in another world. Even if we try to extend the search for 'hard' Darwinians beyond the names on Allen's list, remarkably few candidates present themselves. The attitude of T.H. Huxley, 'Darwin's bulldog', to the natural selection hypothesis may well be guessed from his reluctance to support it at all, either publicly or privately. Though he sprang to align himself with the evolutionists, and though he spoke of his readiness 'to go to the stake, if requisite', in support of the evolutionary thesis per se,⁷ he was cool enough about natural selection in a lecture of 1860 for Darwin to complain about it to their mutual friend Hooker; and twenty years later he managed the uncanny feat of delivering an address celebrating the anniversary of the Origin without mentioning its central postulate at all! If he never let himself be branded an apostate, Huxley was never a disciple either.

We are obliged to conclude, therefore, that the reaction against Darwinism in the first decades after the Origin was a much more profound and firmly reasoned reaction (though indeed highly charged emotionally) than may be apparent at first sight. Scientific history, no less than political, tends to be the victor's version; what is more, as Howard Gruber has reminded us, in the history of ideas it is often the persecutors themselves, the vociferous opponents of a new theory, who go down into silence, with the traces they leave behind on that theory's proponents being quickly effaced by time and historians' indifference.⁸ And here the matter is further complicated by the undoubted fact, conceded by all parties, that one part of Darwin's work -- namely its validation of evolution -- did indeed gain recognition and approval, and at a speed which his great peers Marx and Freud might well have envied. But on inspection post-Darwinian biology is found to relate to the insights of its founder in a much more ambiguous way than do Marxian economics and Freudian psychology. The remainder of this chapter will try to expand on this assertion, taking as its text C.D. Darlington's admittedly somewhat heterodox interpretation that in the period under review Darwinism 'began as a theory that evolution could be explained by natural selection. It ended as a theory that evolution could be explained just as you would like it to be explained.'⁹

Darwinism in Decay, 1870-1900
From our perspective it is difficult to avoid seeing the intellectual history of the 1850s as one inexorable wave sweeping forward with gathering momentum, gathering up all the pre-Darwinian evolutionists into one grand synthesis, and finally breaking with a crash in 1859. We smile to read Darwin's denial that the subject of the origin of species was at all in the air when he was collecting his evidence. And yet there is a sense in which the Darwinian revolution was more of a counter-revolution: in an important way it subverted the persisting Romantic impulse towards progressive evolutionism, which had already demonstrated its power by making Robert Chambers's Vestiges of the Natural History of Creation (1844) a bestseller on a scale far outstripping the Origin's immediate success a decade later. Despite Chambers's scandalous handling of Genesis, the ideas of his book were assimilated, even by the Established Church, much more easily than Darwinian natural selection. Right up to 1859, with astronomy and geology between them producing more and more cheering evidence for cosmic progressivism on the Spencerian model, the church's energies were well absorbed by the Oxford Movement, leaving little to spare for any direct confrontation with science. Samuel Butler grew to maturity in these years of somnolent calm and, as he tells us in his account of the childhood of his alter ego Ernest Pontilex, there was between 1844 and 1859 'not a single book published in England that caused serious commotion within the bosom of the Church'.¹⁰ Even In Memoriam, whose biological assumptions as developed through the whole poem seem to be based at least in part on Chambers, does not constitute a clear exception to the view that the century's fifth decade was a period of quietly optimistic quiescence, marked symbolically at its beginning by the death of Wordsworth and Wordsworthian pantheism and at its end by the publication of The Origin of Species. There is little evidence, even when we take into account the writings of earlier evolutionists like William Wells (1757-1817), Patrick Matthew (1790-1864), William Lawrence (1783-1867) and Edward Blyth (1810-73), of the coming storm; and there is no sign of it at all in the serene poeticised geology of Martin Tupper's Proverbial Philosophy (1838-76) or Kingsley's Glaucus (1855). The cracks in pre-Darwinian evolutionism did not begin to gape really wide until Philip Gosse refused both to regard any part of Scripture as allegorical and to reject the evidence of his natural history researches. His Omphalos (1857), written, in Gibbon's phrase, 'to reconcile the jarring interests of reason and peity', was ridiculed by all parties precisely because it was too commonsensical in attempting to marry jarring Genesis and geology.

In such a climate of opinion that which was least original in the Origin -- the proposal of organic evolution as a historical process -- gained the speediest assent. Darwin did not invent that theory and did not make it familiar to anyone with even a smattering of the geological knowledge of the first half of the century. By the 1840s the transmutation of species had become a serious and established hypothesis; even earlier, in the 1830s, there was circulating a 'bestial theory' of man's affinity with the higher apes -- circulating widely enough for Coleridge to mock it in 1833. In this sense, the Origin supplied no more than a shock of recognition. But the addition of natural selection to supply a propulsive force to evolution was truly revolutionary -- though not in truth absolutely original, for Darwin was anticipated in some degree by all the evolutionists mentioned above, and by others. It was this addition, this unalloyed Darwinism, which sounded through the literary culture as perplexity, outrage or buoyant inspiration. To some extent the proof of this assertion must be a continuous one; but we will pause here to refer briefly to an invaluable semi-statistical survey by Alvar Ellegard which permits us to bring a little much-needed rigour into this particular area of the history of ideas. On the question of the differing rates of diffusion through Victorian culture of belief in the unit-ideas of 'species-transformation' and 'natural selection' Ellegard is most informative. Ten years after the Origin general belief in the evolution (i.e. the progressive transformation) of species ran through every sector of society except, perhaps, a small group of rigidly fundamentalist churchmen. The Darwinian hypothesis, however, in Ellegard's considered opinion, 'clearly met with an incomparably stronger resistance... It touched the ideology of the age at a more vital point than did the Evolution theory pure and simple.'¹¹ Whereas, he contends, the controversy over evolution itself was effectively dead within ten years, Darwin's explanation for it steadily declined in popularity. Admittedly, this judgement is offered with the proviso that in the case of human evolution only a small proportion of the authors surveyed were prepared to accept a totally naturalistic explanation for the origin of man, and particularly for the origin of his higher mental faculties. Certainly on what evidence was available this was a good deal to swallow. But the exception Ellegard makes is not a crucial one. The late-Victorian teleologists did not dissent from the evolutionistic orthodoxy of human functions having developed steadily and progressively; they insisted only that the process must have been guided, and so to a large extent the proponents of that highly influential stream of thought still fit into Ellegard's pattern. But he goes farther than this. His careful study of a wide range of periodical articles devoted to these topics leads him to distinguish two fairly distinct phases in the decline of Darwinism. Among the informed and well-disposed, evolution by natural selection made some headway between 1864 and 1869, for the organised technical opposition to it took some time to become fully mobilised. Six years passed before G.M. Humphry's first elaborate critique was placed before the British Association. During this period the hysterical and highly irrational denunciations of Darwin caused a ground swell of support for him among the scientific popularisers and literary men with fewer or no religious convictions to be outraged. However, in the second period up to 1872 the adherents of Darwinism in its 'hard' or classical form fell away sharply. Ellegard's study unfortunately ends in that year; but since the strictly scientific opposition to Darwinism grew steadily stronger after 1872, and was recognised as becoming more and more formidable by all parties, there is every reason to suppose that a statistical inquiry would show this decline continuing at an accelerating pace up to the end of the century and for some time beyond it. By the middle 1890s a small countervailing tendency first becomes detectable in the rise ofneo-Darwinism; and it is this that marks the end, effectively, of the curious period of suspension that we are trying to define. In modern biology the significance is well recognised of a phenomenon called paedomorphosis, whereby a species can, in order to escape a developmental dead end, regress to an infantile stage in its lifecycle before pushing forward again. The history ofneo-Darwinism is just such an example of a reculer pour mieux sauter which quite reversed the normal pattern of acceptance, consolidation and a slow decline in explanatory power. It was the genius of the neo-Darwinians, and a little later of the first post-Mendelian geneticists, that they were ready and able to backtrack mentally to 1859 and pick up the one possibility which had been generally neglected: that of eliminating entirely from a species-forming process everything except individual differences, the random bursts of mutation and the consequent modifications induced by selection. This was a bold and unpopular move for some time, because even at that late date it was still violently throwing into reverse gear the main drive of evolutionary thought. The evolution of a science, then, like that of a species, need not proceed smoothly. Evolutionary biology is of course now Darwinian, and it is highly unlikely that it will ever move backwards to embrace modes of explanation now thoroughly discarded. A parallel is to be found in the history of physics: quantum mechanics has there replaced the Newtonian-Galilean model, but the net result has been a shift even farther away, conceptually, from scholastic physics. Such is the trend of a science in the long term; but there can be small fluctuations which run for a while in the opposite direction, and it is inside such a fluctuation that our present attention is directed.

So far our chief emphasis has been placed on the emotional, aesthetic and theological causes for this interregnum in biological explanation. And this is correct. For, despite the title of his great work, Darwin did not believe that species really exist apart from the observer. A philosophical nominalist like Lamarck before him and Vries after, Darwin spoke of species as a mere taxonomic convenience. For in truth it is not species in the abstract which bear the brunt of selection, but unique organisms whose minute differences from each other are so readily overlooked for the benefit of the systematist. Yet it was just these minutiae, and not the intraspecies similarities, that Darwinism in its pre-Mendelian phase stressed as supremely important, for it is these minutiae alone which are the material of evolutionary change. Natural selection therefore cut the ground from under not merely the theological arguments from design, but even more disruptively from under the deep-seated belief in the order, economy and regularity of nature. Out of the Enlightenment, and especially out of its astronomy, had come the apparent discovery that there is a basic rationality in the world order a rationality finding an encouraging reflection in the a priori axioms of mathematics. One could juggle one's indisputably home-made symbols; one could come up with a set of co-ordinates (that is, if one were a Le Verrier or an Adams); a telescope could be directed to a precise point in the sky, and a new planet (Neptune) could sweep into one's ken. Between the inner workings of the human brain and the majestic sweep of the planets a most pleasing concordance seemed to prevail. Darwinism changed all that. The strongly cohesive bonds which held men together in the dignity of Homo sapiens lost their grip. Now there were only individual men, more or less wise as the caprice of variation would have it. And how could a natural law whose very hallmark ought to be lucidity, beauty and simplicity work by generating a thousand, a million more waste products than perfected items? Such, at least, is the logic of one popular train of argument. There were others.

This being said, it is important as a corrective to make mention of the very severe difficulties of a strictly technical kind which assailed the Darwinian theory and made it quite possible for anyone who was cold to it to decline to take it seriously. Darwin did his best to make a completely natural science of biology as a whole, to outlaw teleology and to substitute detectable changes happening lawfully over geological ages; but on any analysis regularity and observability were not strong points of the mechanism Darwin posited. Variation followed no law known to man and no one, certainly not Darwin himself, could make any claim to have watched species forming in the wild. Nor did he receive the support on which he had relied from the neighbouring sciences. The enunciation of the basic laws of genetics lay forty years ahead. Palaeontology, despite Lyell's valiant support for his friend and colleague, could not demonstrate the mutability of species in the past but only that the more recent species tend to a greater complexity of function. In embryology and its puzzling datum that ontology repeats phylogeny in utero Darwin believed he had an ally; but, though recapitulation was well understood by Baer and Agassiz, these formidable men nevertheless rejected it as evidence of evolution. And biochemistry was at a complete loss to explain the conversion of the same primitive embryonic tissue into quite distinct organs. 'What directive agency determines the distribution of the colouring matter,' Wallace was still pertinently asking of embryology and biochemistry a decade after the close of our period as he had asked for the whole span of it, 'so that each feather shall take its exact share in the production of the whole pattern and colouring of the bird, which is immensely varied, yet always symmetrical as a whole, and has always a purpose'?¹² To this and allied questions, which step far beyond the limits of evolutionary biology as it was then studied, Darwinism had in the first four decades of its existence no clear and distinct reply.

Still less could Darwinism cope with a challenge from outside biology altogether. No attack on the foundations of the Darwinian edifice gained greater credence than that by the physicist William Thomson, later Lord Kelvin (1824-1907), who came to biological controversy with the fame of his brilliant formulation of the Second Law of Thermodynamics (1851) securely behind him. Overtly at least, Thomson never had much interest in the details of species modification or indeed in the life sciences at all. He restricted himself to arcane problems of geology, mathematics and astrophysics, and he became so challenging an opponent of the Darwinians because he claimed to undercut their position on physical evidence alone.

In the first edition of The Origin of Species Darwin laid very great stress on the nearly unimaginable scale of geologic history; on the passage of hundreds of millions of years during which time, if the Uniformitarians were correct, conditions for life had been much as they are today. Sheer time had for Darwin a creative aspect. Was there anything, no matter how improbable, that could not have recurred time and time again somewhere in that vast bulk of years? Could not even natural selection, running its slow and fumbling course, achieve anything, even man himself, in 300 million to 500 million years? Here Darwin had a convincing argument and one which he deployed with considerable skill. Instead of speaking of aeons of time in the abstract he brings before our mind's eye an actual example of sedimentary formations nearly fourteen miles thick; then, instead of supplying us gratis with an estimate of how long it might have taken this to form, he tells us that the mighty Mississippi lays down about 600 feet of sediment in a hundred thousand years (six feet per thousand years has a far less magnificent thud to it). He leaves the further impressive calculation to us. When we connect up this with the fact that whole generations of simple organisms may succeed each other in days or even in hours, this deftly handled ninth chapter of the Origin on the geological record becomes the keystone to the book. Man, too, had his place in this geological record. Charles LyeIl (1797-1875), the most famous geologist of the day, had rejected transformism in his Principles of Geology (1830-3). The effect, then, was all the greater when in his Antiquity of Man (1863) he not only faced about but did Darwin the service of pushing back many thousands of years the emergence of Homo sapiens and his predecessors, citing new palaeontological evidence for so doing. His reputation helped -- but for a while only -- to suggest that geology and Darwinism were in harmony.

But Thomson asked a simple question and, in doing so, cooly sliced away the Darwinians' vast womb of time. If all life depends on the sun's heat, how old is the sun? In forming his estimate it goes without saying that Thomson had no inkling of the Bethe reaction, the thermonuclear conversion process (not described until 1939) which sustains the sun's colossal energy output. He could only imagine a mechanical source of heat, or a chemical one. In a typical early paper he dismisses such sources of energy as meteors failing into the sun or the sun's contracting under its own gravitational attraction. He is led to assume that the sun is one huge glowing coal, a mere 'incandescent liquid mass cooling'.¹³ From there, given the known properties of combustible materials, it was a relatively trivial calculation to come up with the judicious statement that 'all geological history showing continuity of life, must be limited within some such period of past time as one hundred million years'.¹⁴ Victorian astrophysics was not a quickly moving science. Nineteen years later Thomson had few additional facts to offer, but those he did have made him slash his allowance to the biologists down to a mere 20 million years¹⁵ -- a time now known to have covered the emergence of only the most recent mammals. Thomson continued to insist on this alarmingly small figure right down to the end of the century.¹⁶

The consternation within the Darwinian circle produced by Thomson's calculations was immense, and it certainly assisted materially in the decline of Darwinism from the early 1870s. The alarm, however, grew slowly. None of the Darwinians gave any public recognition of the deadly significance of Thomson's work before 1868 at the earliest, though they did so privately. The first four editions of The Origin of Species (ie from 1859 to 1866) do not mention Thomson by name, although as early as the second edition the time mentioned as being available for evolution to work in was scaled down considerably. Thomson could be ignored: he had been cautious in making his foray into alien territory and perhaps had muted the inevitable biological conclusions in the hope that others would make the necessary connections. Meanwhile, the Darwinians' attentions were fully occupied by the immediate threat posed by the most carefully worked-out of the early attacks -- that of Fleeming Jenkin. The burden of Jenkin's mathematical analysis of 1867 was that, even if a favourable variation should occur by chance in an individual, the likelihood of its being spread through the population was remote unless both parents displayed it. Otherwise the useful novelty would be diluted Clenkin wrongly took the blending of characteristics for granted) and lost like a drop of ink in the ocean. Faced with these relatively straightforward puzzles the Darwinians and their public could pass over the unworldly abstractions of Thomson's cosmogonical theories. But, whatever the reason for neglect, Thomson threw down the gauntlet unmistakably during his lecture before the Geological Society of Glasgow in February 1868. The traumatic effect on Darwin personally left many traces in his writings. A little more than a year after this lecture Thomson had become one of Darwin's 'sorest troubles' and two years after this 'an odious spectre' -- strong language for the mild Darwin, even to an intimate.¹⁷ In the spring of 1869, while revising the Origin for its fifth edition, Darwin privately summarised one important concession that he was about to make rather misleadingly as '1 have been led ... to think the lapse of time, as measured by years, not quite so great as most geologists have thought',¹⁸ whereas in the forthcoming edition he was in fact going to curtail his estimate of the epochs available for the evolution of life hardly to one 'not quite so great' as before but to a figure less than one-third the size of his original. But this was in a private letter: in public Thomson received more justice than one might have expected. When the fifth edition came out (1869) we hear of his 'formidable objection',¹⁹ and by the sixth (1872) this has been strengthened to 'probably one of the gravest as yet advanced' (XIV:55.l:f) and one detects also a rather desperate quality in the countering arguments. When we set Darwin's final words that 'we do not know at what rate species change as measured by years' (XIV: 55.1f) against his earlier assurance that 'an infinite number of generations' (IX:74) have undergone almost infinitely slow change, we have the most direct evidence of the destructive power of Thomson's papers. A particular irony of the situation was that every increase in the known length of man's prehistory made the problem that much more acute. As man and his prehuman ancestors were traced back more than a million years, how could the whole succession of life from amoeba to man be jammed into an allowance of years only some twenty times longer? Most of the Darwinians eventually broke down at this point: natural selection could not have worked within such a period. Only some teleological mechanism from within, or some transcendent will from without, could have worked speedily enough. Others retreated into sophistries. T.H. Huxley's lecture 'Geological Reform' of 1869 in particular marks a very low point indeed in that skilled debater's powers to muster an irrefragable argument. Despite Darwin's own stress on the capacity of slowly passing time to build on random and irregular variations Huxley does his best to convince us that

biology takes her time from geology. The only reason we have for believing in the slow rate of the change in living forms is the fact that they persist through a series of deposits which, geology inform us, have taken a long while to make. If the geological clock is wrong, all the naturalist will have to do is to modify his notions of the rapidity of change accordingly.²⁰

The bland speciousness of this is alarming. It is a characteristic of Huxley's rhetoric that it becomes more facile as the insecurity of his stance grows; and by this measure Thomson had made him very insecure indeed. The rate of change of living structures was no mathematical variable to be stretched or chopped to fit the Procrustean bed Thomson had prepared for it. Either that change had come about thanks to the Darwinian mechanism assisted by time, or there was some other vera causa of evolution -- if evolution there had been. Thomson's reply a month or so later is a model of amiable butchery. Joint by joint Huxley's feeble defence is dismembered. The Darwinian mechanism will only work where there is a wealth of slow time. Cut down the allowance by a factor of ten or more and the conclusion is inevitable:

the limitation of geological periods, imposed by physical science, cannot, of course, disprove the hypothesis of transmutation of species; but it does seem sufficient to disprove the doctrine that transmutation has taken place through 'descent with modification by natural selection'.²¹

Evolution is a fact, but something, or someone, has guided its path. If life had had to wait on some 'fortuitous concourse of atoms', then there would even now be nothing but a wash of protein molecules in a dead sea. As far as Thomson, most other geophysicists and an ever-increasing number of laymen were concerned, that was that for the remainder of the century. Thomson happily reprinted his lectures in 1889 and 1894 without altering a word.

In 1903 the discovery of Curie and others that the element radium has a temperature permanently above the ambient level wrecked Thomson's elegant structure, for it was then only necessary to suppose that radioactive elements are present in the sun for Thomson's estimates to be multiplied by any figure one wished. Geological time expanded again so rapidly that no Edwardian physicist dared to offer any calculation as to either the probable or possible age of life on earth, especially after the lead-uranium technique for dating rocks began to give figures in the billion- rather than the hundred-million-year range. The twistings and turnings of the Darwinians proved in the end to have been unnecessary. For several decades, however, the physical sciences seemed linked in a reactionary conspiracy with the fundamentalists to make the creation recorded in Genesis seem less of an outrage on common sense than that proposed by the new biology. It followed that one surrendered no intellectual respectability by refusing to be a Darwinian.

Darwinism reached the nadir of its reputation in the three decades round the turn of the century. It had been abused from all sides, replaced time and again by some version of evolutionism and abandoned by one after another of its advocates including to quite some extent even Darwin himself before his death. It is only the first edition of The Origin of Species that has very much argumentative consistency. Later editions became palimpsests of qualification and contradiction as Darwin struggled with his major critics, and the fifth and sixth editions contain more than half of the total number of revisions made by the author in his lifetime. Nor were these revisions well received. Darwin himself mentioned in a letter to Hooker of July 1868 that he had heard 'the belief in Natural Selection is passing away'. Failing to placate his critics, and perhaps knowing full well that some of the criticisms were irrefutable within the prevailing limits of knowledge, Darwin retreated in the last decade of his life to more neutral ground. In treating of the habits of earthworms (1881) he could apply his unmatched gifts for extended and minute observation to the investigation of such matters as the rate of burial of ancient buildings, without having to keep one eye all the time on treacherous definitions which seemed to grow steadily more unruly as the years passed.

Darwin did not retreat, though, without making heroic efforts to cope -- efforts which unfortunately compelled him to give full rein to the more credulous side of his nature. Even in the first edition of The Origin of Species he neither wished nor was able to dispose entirely of the evidence for the inheritance of acquired characteristics. Despite the taint of teleology Darwin held on to at least two neo-Lamarckian axioms. He allowed from the start that certain variations are not random but must be due either to changes induced by the environment or to the effects of use or disuse on organs or habits, or to both. Nor did he hold out for very long against even Lamarck's notion of a sentiment interieur in each creature; but these constituent parts of the original Lamarckian system tend to become very muddled in Victorian neo-Lamarckism and for present purposes it is not necessary to distinguish them. We need only to notice how under extreme critical pressure several varieties of post-Darwinian speculation allowed purposiveness, design and the concept of a life-force to creep back through the holes in the argument of the Origin -- an argument in its naked form genuinely proof against all non-naturalism. This is well exemplified in Darwin's own evaluation of Lamarckism, which moves in a full circle from elaborate praise in the early notebooks, through a verdict of 1863 on the Philosophie zoologique as being 'a wretched book, and one from which (I well remember my surprise) I gained nothing',²² on to a renewal of Lamarckian explanation in the The Variation of Animals and Plants under Domestication (1868), and at last back to the addition of a whole new chapter describing goal-directed variation in the sixth edition of The Origin of Species (1872). In this ambiguous chapter Darwin now appeals in fairly equal proportions to chance (unknown law) and to the conditions of life as the true causes of variation, though to deal effectively with Thomson on his own terms meant that much stress had to be placed on the idea of environmental factors somehow speedily compelling a required organ into existence. But, whatever the pressures on Darwin as a theoreticjan, looking at his new invocation of Lamarckism more broadly obliges us to see in it a reflection of the very pronounced desire among philosophers, social critics and educationalists to believe that acquired characters are transmissible down the generations. In such new guise did the progressivism of the 1840s and 1850s turn up again. Herbert Spencer's adaptive sociology, spanning the pre- and post-Darwinian eras, was rooted in the innate psychology of Gall, but to make his sociology truly evolutionistic after 1859 Spencer had to give a sovereign place to Lamarckian inheritance. Indeed, his reliance on it was so great that to abandon it would have meant abandoning his whole philosophy. Fortunately he was not called upon to do so since the evidence was still ambiguous at the date of his death: it became clear only very much later; and even then, as Zirkle records of the Lamarckians' last gasp in Soviet genetics, 'perhaps no other biological hypothesis has ever been tested so thoroughly and abandoned with such reluctance'.²³ Some of the untenable generalisations in The Variation of Animals and Plants sound as if they are cultural reflections, as when Darwin quotes solemnly from Saint-Hilaire that 'the mothers of illegitimate children troubled in their minds and forced to conceal their state, are far more liable to give birth to monsters than women in easy circumstances'.²⁴ There may be a grain of physiological truth in this, but what Darwin is surely trying to implant is the suspicion that bastards are the unfit, to be weeded out by a selection which is not merely neutral and natural but social too, and in favour of legitimate offspring.

Apart from this kind of transparent response to cultural pressures, the Darwinians made increasing appeals to teleology because it helped to make intelligible the curiously spasmodic irregularities in the frequency of variation. The common supposition of the time was that scientific methodology must rise from the bottom-most rung of explanation if it was to avoid the charge of arguing ignotum per ignotius. Certainly Darwin's handling of variation was clumsy; and certainly while Mendelism lay hid in night the ghost of teleology could not be laid. While the lacuna existed there was a constant pressure to fill it up with some directive, and usually divine, process. The American Asa Gray was ready almost as soon as the Origin was off the press with the analogy of an irrigation stream. The beds of such streams, he argued, are formed in exactly the same way as the beds of the natural rivers. The water, according to the fixed and impersonal law of gravity, flows to the lowest point and in doing so rolls down stones which wear out a channel by the equally fixed laws of friction. The designer of an irrigation ditch does not try to flout these laws. He limits himself to directing the flow in the required direction. May we not believe, therefore, that, while natural selection orders the details, the evolutionary flow as a whole is the result of the Divine Irrigationist forcing the stream of variation down one channel rather than another? Indeed, Gray insisted, it would be gross scientific irresponsibility 'at least while the physical cause of variation is utterly unknown and mysterious' not to believe that 'variation has been led along certain beneficial lines'.²⁵ Nature should be assumed innocent and supportive until proven uncaring and random.

Darwin's own musings on the broader implications of his theory were rarely more sophisticated than Gray's, but (in private at least) he concurred with Gray that his 'chance' as the cause of variation would not do -- it too easily becomes a wastepaper-basket to receive any inconvenient difficulties. When he wrote to Asa Gray that the prime mover in his theory was random variation, he speedily added the rider 'not that this notion at all satisfies me. I feel most deeply that the whole subject is too profound for the human intellect. A dog might as well speculate on the mind of Newton.'²⁶ The odd touch of philosophic idealism in that last sentence (the hint that the organic universe may be more like a great thought than a great roulette-wheel) is indicative of that perennial inclination in Darwinian theory: the tendency to drift into using internal or external purposiveness as a catch-all unless firmly restrained from doing so. In The Descent of Man that drift is almost complete. By this time (1871) Darwin would probably have been content, had it not been for the impetus supplied by his public's expectations, to leave the task of elucidating man's own evolution by natural selection to others. We know that he did in fact attempt to pass his notes for the Descent on to Wallace, hoping that he would shoulder the enormous burden; but sensibly Wallace declined, almost certainly out of a conviction that it could not be done. It is, then, not very surprising to find that the Descent lacks the solid drive of a single sustained argument so characteristic of the early Origin. Instead it gives the impression of having grown by accretion, of having fact piled upon fact until it is overweighted with information. And there is a discomfiting toleration of all sources of this information as having equal merit, so that anecdotes from proud pet-owners mingle confusingly with the matter-of-fact expertise of stockbreeders. The refusal to accept variation as random and uncaused is carried to the point where we are told that Europeans in America undergo after a few generations 'a slight but extraordinary [sic] rapid change of appearance. Their bodies and limbs become elongated.'²⁷ Conceptually we are here miles away from that careful experimenter who formerly had weighed duck skeletons and sought for viable seeds in the excrement of seabirds.

Anti-Darwinian Thought Concluded
The purpose of this all-too-rapid excursion through what is still an imperfectly understood sector of scientific history will have been amply fulfilled if it has brought home just how divergent opinion was, even among the most detached theoreticians, of the human significance of evolutionary biology. The opinionative spectrum ran all the way from Spencer's influential belief that progress (moral and spiritual) is implicit within the evolutionary progression from spiral nebulae to the refinements of Victorian society and beyond; to the aged Huxley's equally influential belief that cosmic progress is fundamentally amoral and must be vigorously opposed and rejected for civilised life to be sustained and improved. No compromise is possible here, and we should not seek to impose one. These divisions emerged out of a particularly static phase in scientific history, and with their relative truth or falsity we are not concerned. When, in the following chapters, there is a charting of the sometimes extreme manipulations of biological concepts it should be borne in mind that there was no reason in the world why even the best-informed novelist or the most cautious speculative essayist, each in his own way concerned to present the latest truths of the life sciences in all their purity, should have adhered to any orthodox line. Natural selection, to mention only the key item, was certainly a bridge which led from romantic pantheism and natural theology. But it led to no clear destination. Positivism, chiliasm, reworked pantheism, neoPaleyism, stoical despair, guarded optimism, even mystical exaltation: there were expert defendants for all these and more. None of these conflicting pictures of the world alone represents the supreme contribution of Victorian biology to cultural history.

Our excursion may have given the impression that there was little more for the literary imagination to seize on in Victorian biology than the conflict over randomness versus teleology in variation and inheritance. An important conflict it certainly was, equally for the philosopher concerned for progress, for the reformer intent on social change, and for the moralist worried about the inheritance of virtue and vice. These professional concerns all left a mark on literature. But biology also engendered other equally profound shifts in orientation which were just as amenable to imaginative exploitation. There was the concept of a single primitive substance, protoplasm, as the basic material of all life from bacteria to man. There was the discovery that fundamental metabolic processes work on the same principles in all life-forms, even in those most dissimilar to the human. There was the growing tendency in all the life sciences to reductionism -- first to accounting for phenotype in terms of genotype, but later more drastically to the mechanistic assumptions of the contemporary physics. Vitalism was exploded by Wohler's synthesis of urea in 1828, and something which could almost be taken for 'growth' was displayed in inorganic crystals by Schleiden in 1838. The distinction between the organic and the inorganic was thereby shown to be no more than a convenience of terminology. Organisms were of the same substance through and through as the inanimate cosmos, and were subject to all the laws of a universe made out of Daltonjan atoms: unchanging, unyielding billiard-balls bounced from one configuration to another by physico-chemical forces. Biology was no sooner thoroughly entangled in this inadequate metaphysical net than developments in the physical sciences began to rip it apart; but while the model lasted it forced biology into a series of antitheses (mind or matter, determinism or randomness, Darwinism or Lamarckism) from which no writer of scientific inclinations could remain aloof. Finally and most crucially there was the common (though not universal) ejection from the new theories of evolution of the ancient doctrine of the chain of being, together with all its inbuilt hierarchical suppositions. To certain mentalities man's image of himself as an intermediate link in this chain subsided into a new picture of a many-branched tree where the branch occupied at present by man might yet fork again, bringing something new, superior and alien. All these conceptions (and they were equally useful whether accepted or denied) were fraught with literary possibilities. Before examining some of these in detail, we need some account of the part played by biology in the extra-scientific culture, and we shall approach this by considering some of the contemporary attitudes to science as a fertilising force on literature.