Biogenesis And Abiogenesis

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Title:     Biogenesis And Abiogenesis
Author: Thomas Henry Huxley [More Titles by Huxley]

(THE PRESIDENTIAL ADDRESS TO THE BRITISH ASSOCIATION FOR THE ADVANCEMENT OF SCIENCE FOR 1870)

It has long been the custom for the newly installed President of the British Association for the Advancement of Science to take advantage of the elevation of the position in which the suffrages of his colleagues had, for the time, placed him, and, casting his eyes around the horizon of the scientific world, to report to them what could be seen from his watch-tower; in what directions the multitudinous divisions of the noble army of the improvers of natural knowledge were marching; what important strongholds of the great enemy of us all, ignorance, had been recently captured; and, also, with due impartiality, to mark where the advanced posts of science had been driven in, or a long-continued siege had made no progress.

I propose to endeavour to follow this ancient precedent, in a manner suited to the limitations of my knowledge and of my capacity. I shall not presume to attempt a panoramic survey of the world of science, nor even to give a sketch of what is doing in the one great province of biology, with some portions of which my ordinary occupations render me familiar. But I shall endeavour to put before you the history of the rise and progress of a single biological doctrine; and I shall try to give some notion of the fruits, both intellectual and practical, which we owe, directly or indirectly, to the working out, by seven generations of patient and laborious investigators, of the thought which arose, more than two centuries ago, in the mind of a sagacious and observant Italian naturalist.

It is a matter of everyday experience that it is difficult to prevent many articles of food from becoming covered with mould; that fruit, sound enough to all appearance, often contains grubs at the core; that meat, left to itself in the air, is apt to putrefy and swarm with maggots. Even ordinary water, if allowed to stand in an open vessel, sooner or later becomes turbid and full of living matter.

The philosophers of antiquity, interrogated as to the cause of these phenomena, were provided with a ready and a plausible answer. It did not enter their minds even to doubt that these low forms of life were generated in the matters in which they made their appearance. Lucretius, who had drunk deeper of the scientific spirit than any poet of ancient or modern times except Goethe, intends to speak as a philosopher, rather than as a poet, when he writes that "with good reason the earth has gotten the name of mother, since all things are produced out of the earth. And many living creatures, even now, spring out of the earth, taking form by the rains and the heat of the sun."[1] The axiom of ancient science, "that the corruption of one thing is the birth of another," had its popular embodiment in the notion that a seed dies before the young plant springs from it; a belief so widespread and so fixed, that Saint Paul appeals to it in one of the most splendid outbursts of his fervid eloquence:--

"Thou fool, that which thou sowest is not quickened, except it die."[2]

[Footnote 1: It is thus that Mr. Munro renders

"Linquitur, ut merito maternum nomen adepta
Terra sit, e terra quoniam sunt cuncta creata.
Multaque nunc etiam exsistant animalia terris
Imbribus et calido solis concreta vapore."

_De Rerum Natura_, lib. v. 793-796.

But would not the meaning of the last line be better rendered "Developed in rain-water and in the warm vapours raised by the sun"?]

[Footnote 2: 1 Corinthians xv. 36.]

The proposition that life may, and does, proceed from that which has no life, then, was held alike by the philosophers, the poets, and the people, of the most enlightened nations, eighteen hundred years ago; and it remained the accepted doctrine of learned and unlearned Europe, through the Middle Ages, down even to the seventeenth century.

It is commonly counted among the many merits of our great countryman, Harvey, that he was the first to declare the opposition of fact to venerable authority in this, as in other matters; but I can discover no justification for this widespread notion. After careful search through the "Exercitationes de Generatione," the most that appears clear to me is, that Harvey believed all animals and plants to spring from what he terms a "_primordium vegetale_," a phrase which may nowadays be rendered "a vegetative germ"; and this, he says, is _"oviforme_," or "egg-like"; not, he is careful to add, that it necessarily has the shape of an egg, but because it has the constitution and nature of one. That this "_primordium oviforme_" must needs, in all cases, proceed from a living parent is nowhere expressly maintained by Harvey, though such an opinion may be thought to be implied in one or two passages; while, on the other hand, he does, more than once, use language which is consistent only with a full belief in spontaneous or equivocal generation.[3] In fact, the main concern of Harvey's wonderful little treatise is not with generation, in the physiological sense, at all, but with development; and his great object is the establishment of the doctrine of epigenesis.

[Footnote 3: See the following passage in Exercitatio I.:--"Item _sponte nascentia_ dicuntur; non quod ex _putredine_ oriunda sint, sed quod casu, naturae sponte, et aequivoca (ut aiunt) generatione, a parentibus sui dissimilibus proveniant." Again, in _De Uteri Membranis:_--"In cunctorum viventium generatione (sicut diximus) hoc solenne est, ut ortum ducunt a _primordio_ aliquo, quod tum materiam tum elficiendi potestatem in se habet: sitque, adeo id, ex quo et a quo quicquid nascitur, ortum suum ducat. Tale primordium in animalibus (_sive ab aliis generantibus proveniant, sive sponte, aut ex putredine nascentur_) est humor in tunica, aliquaaut putami ne conclusus." Compare also what Redi has to say respecting Harvey's opinions, _Esperienze_, p. 11.]

The first distinct enunciation of the hypothesis that all living matter has sprung from pre-existing living matter, came from a contemporary, though a junior, of Harvey, a native of that country, fertile in men great in all departments of human activity, which was to intellectual Europe, in the sixteenth and seventeenth centuries, what Germany is in the nineteenth. It was in Italy, and from Italian teachers, that Harvey received the most important part of his scientific education. And it was a student trained in the same schools, Francesco Redi--a man of the widest knowledge and most versatile abilities, distinguished alike as scholar, poet, physician, and naturalist--who, just two hundred and two years ago, published his "Esperienze intorno alla Generazione degl' Insetti," and gave to the world the idea, the growth of which it is my purpose to trace. Redi's book went through five editions in twenty years; and the extreme simplicity of his experiments, and the clearness of his arguments, gained for his views, and for their consequences, almost universal acceptance.

Redi did not trouble himself much with speculative considerations, but attacked particular cases of what was supposed to be "spontaneous generation" experimentally. Here are dead animals, or pieces of meat, says he; I expose them to the air in hot weather, and in a few days they swarm with maggots. You tell me that these are generated in the dead flesh; but if I put similar bodies, while quite fresh, into a jar, and tie some fine gauze over the top of the jar, not a maggot makes its appearance, while the dead substances, nevertheless, putrefy just in the same way as before. It is obvious, therefore, that the maggots are not generated by the corruption of the meat; and that the cause of their formation must be a something which is kept away by gauze. But gauze will not keep away aeriform bodies, or fluids. This something must, therefore, exist in the form of solid particles too big to get through the gauze. Nor is one long left in doubt what these solid particles are; for the blowflies, attracted by the odour of the meat, swarm round the vessel, and, urged by a powerful but in this case misleading instinct, lay eggs out of which maggots are immediately hatched, upon the gauze. The conclusion, therefore, is unavoidable; the maggots are not generated by the meat, but the eggs which give rise to them are brought through the air by the flies.

These experiments seem almost childishly simple, and one wonders how it was that no one ever thought of them before. Simple as they are, however, they are worthy of the most careful study, for every piece of experimental work since done, in regard to this subject, has been shaped upon the model furnished by the Italian philosopher. As the results of his experiments were the same, however varied the nature of the materials he used, it is not wonderful that there arose in Redi's mind a presumption, that, in all such cases of the seeming production of life from dead matter, the real explanation was the introduction of living germs from without into that dead matter.[4] And thus the hypothesis that living matter always arises by the agency of pre-existing living matter, took definite shape; and had, henceforward, a right to be considered and a claim to be refuted, in each particular case, before the production of living matter in any other way could be admitted by careful reasoners. It will be necessary for me to refer to this hypothesis so frequently, that, to save circumlocution, I shall call it the hypothesis of _Biogenesis_; and I shall term the contrary doctrine--that living matter may be produced by not living matter--the hypothesis of _Abiogenesis_.

[Footnote 4: "Pure contentandomi sempre in questa ed in ciascuna altro cosa, da ciascuno piu savio, la dove io difettuosamente parlassi, esser corretto; non tacero, che per molte osservazioni molti volti da me fatte, mi sento inclinato a credere che la terra, da quelle prime piante, e da quei primi animali in poi, che ella nei primi giorni del mondo produsse per comandemento del sovrano ed omnipotente Fattore, non abbia mai piu prodotto da se medesima ne erba ne albero, ne animale alcuno perfetto o imperfetto che ei se fosse; e che tutto quello, che ne' tempi trapassati e nato e che ora nascere in lei, o da lei veggiamo, venga tutto dalla semenza reale e vera delle piante, e degli animali stessi, i quali col mezzo del proprio seme la loro spezie conservano. E se bene tutto giorno scorghiamo da' cadaveri degli animali, e da tutte quante le maniere dell' erbe, e de' fiori, e dei frutti imputriditi, e corrotti nascere vermi infiniti--

'Nonne vides quaecunque mora, fluidoque calore
Corpora tabescunt in parva animalia verti'--

Io mi sento, dico, inclinato, a credere che tutti quei vermi si generino dal seme paterno; e che le carni, e l' erbe, e l' altre cose tutte putrefatte, o putrefattibili non facciano altra parte, ne abbiano altro ufizio nella generazione degl' insetti, se non d'apprestare un luogo o un nido proporzionato, in cui dagli animali nel tempo della figliatura sieno portati, e partoriti i vermi, o l' uova o l' altre semenze dei vermi, i quali tosto che nati sono, trovano in esso nido un sufficiente alimento abilissimo per nutricarsi: e se in quello non son portate dalle madri queste suddette semenze, niente mai, e replicatamente niente, vi s' ingegneri e nasca."--REDI, _Esperienze_, pp. 14-16.]

In the seventeenth century, as I have said, the latter was the dominant view, sanctioned alike by antiquity and by authority; and it is interesting to observe that Redi did not escape the customary tax upon a discoverer of having to defend himself against the charge of impugning the authority of the Scriptures;[5] for his adversaries declared that the generation of bees from the carcase of a dead lion is affirmed, in the Book of Judges, to have been the origin of the famous riddle with which Samson perplexed the Philistines:--

Out of the eater came forth meat,
And out of the strong came forth sweetness.

[Footnote 5: "Molti, e molti altri ancora vi potrei annoverare, se non fossi chiamato a rispondere alle rampogne di alcuni, che bruscamente mi rammentano cio, che si legge nel capitolo quattordicesimo del sacrosanto Libro de' giudici ... "--REDI, _loc. cit._ p. 45.]

Against all odds, however, Redi, strong with the strength of demonstrable fact, did splendid battle for Biogenesis; but it is remarkable that he held the doctrine in a sense which, if he lead lived in these times, would have infallibly caused him to be classed among the defenders of "spontaneous generation." "Omne vivum ex vivo," "no life without antecedent life," aphoristically sums up Redi's doctrine; but he went no further. It is most remarkable evidence of the philosophic caution and impartiality of his mind, that although he had speculatively anticipated the manner in which grubs really are deposited in fruits and in the galls of plants, he deliberately admits that the evidence is insufficient to bear him out; and he therefore prefers the supposition that they are generated by a modification of the living substance of the plants themselves. Indeed, he regards these vegetable growths as organs, by means of which the plant gives rise to an animal, and looks upon this production of specific animals as the final cause of the galls and of, at any rate, some fruits. And he proposes to explain the occurrence of parasites within the animal body in the same way.[6]

[Footnote 6: The passage (_Esperienze_, p. 129) is worth quoting in full:--

"Se dovessi palesarvi il mio sentimento crederei che i frutti, i legumi, gli alberi e le foglie, in due maniere inverminassero. Una, perche venendo i bachi per di fuora, e cercando l' alimento, col rodere ci aprono la strada, ed arrivano alla piu interna midolla de' frutti e de' legni. L'altra maniera si e, che io per me stimerei, che non fosse gran fatto disdicevole il credere, che quell' anima o quella virtu, la quale genera i fiori ed i frutti nelle piante viventi, sia quella stessa che generi ancora i bachi di esse piante. E chi sa, forse, che molti frutti degli alberi non sieno prodotti, non per un fine primario e principale, ma bensi per un uffizio secondario e servile, destinato alla generazione di que' vermi, servendo a loro in vece di matrice, in cui dimorino un prefisso e determinato tempo; il quale arrivato escan fuora a godere il sole.

"Io m' immagino, che questo mio pensiero non vi parra totalmento un paradosso; mentro farete riflessione a quelle tanto sorte di galle, di gallozzole, di coccole, di ricci, di calici, di cornetti ed i lappole, che son produtte dalle quercel, dalle farnie, da' cerri, da' sugheri, da' leeci e da altri simili alberi de ghianda; imperciocche in quello gallozzole, e particolarmente nelle piu grosse, che si chiamano coronati, ne' ricci capelluti, che ciuffoli da' nostri contadini son detti; nei ricci legnosi del cerro, ne' ricci stellati della quercia, nelle galluzze della foglia del leccio si vede evidentissimamente, che la prima e principale intenzione della natura e formare dentro di quelle un animale volante; vedendosi nel centro della gallozzola un uovo, che col crescere e col maturarsi di essa gallozzola va crescendo e maturando anch' egli, e cresce altresi a suo tempo quel verme, che nell' uovo si racchiude; il qual verme, quando la gallozzola e finita di maturare e che e venuto il termine destinato al suo nascimento, diventa, di verme che era, una mosca.... Io vi confesso ingenuamente, che prima d'aver fatte queste mie esperienze intorno alla generazione degl' insetti mi dava a credere, o per dir meglio sospettava, che forse la gallozzola nascesse, perche arrivando la mosca nel tempo della primavera, e facendo una piccolissima fessura ne' rami piu teneri della quercia, in quella fessura nascondesse uno de suoi semi, il quale fosse cagione che sbocciasse fuora la gallozzola; e che mai non si vedessero galle o gallozzole o ricci o cornetti o calici o coccole, se non in que' rami, ne' quali le mosche avessero depositate le loro semenze; e mi dava ad intendere, che le gallozzole fossero una malattia cagionata nelle querce dalle punture delle mosche, in quella giusa stessa che dalle punture d'altri animaletti simiglievoli veggiamo crescere de' tumori ne' corpi degli animali."]

It is of great importance to apprehend Redi's position rightly; for the lines of thought he laid down for us are those upon which naturalists have been working ever since. Clearly, he held _Biogenesis_ as against _Abiogenesis;_ and I shall immediately proceed, in the first place, to inquire how far subsequent investigation has borne him out in so doing.

But Redi also thought that there were two modes of Biogenesis. By the one method, which is that of common and ordinary occurrence, the living parent gives rise to offspring which passes through the same cycle of changes as itself--like gives rise to like; and this has been termed _Homogenesis_. By the other mode, the living parent was supposed to give rise to offspring which passed through a totally different series of states from those exhibited by the parent, and did not return into the cycle of the parent; this is what ought to be called _Heterogenesis_, the offspring being altogether, and permanently, unlike the parent. The term Heterogenesis, however, has unfortunately been used in a different sense, and M. Milne-Edwards has therefore substituted for it _Xenogenesis_, which means the generation of something foreign. After discussing Redi's hypothesis of universal Biogenesis, then, I shall go on to ask how far the growth of science justifies his other hypothesis of Xenogenesis.

The progress of the hypothesis of Biogenesis was triumphant and unchecked for nearly a century. The application of the microscope to anatomy in the hands of Grew, Leeuwenhoek, Swammerdam, Lyonnet, Vallisnieri, Reaurnur, and other illustrious investigators of nature of that day, displayed such a complexity of organisation in the lowest and minutest forms, and everywhere revealed such a prodigality of provision for their multiplication by germs of one sort or another, that the hypothesis of Abiogenesis began to appear not only untrue, but absurd; and, in the middle of the eighteenth century, when Needham and Buffon took up the question, it was almost universally discredited.[7]

[Footnote 7: Needham, writing in 1750, says:--

"Les naturalistes modernes s'accordent unaninement a etablir, comme une verite certaine, que toute plante vient do sa semence specifique, tout animal d'un oeuf ou de quelque chose d'analogue preexistant dans la plante, ou dans l'animal de meme espece qui l'a produit."--_Nouvelles Observations_, p. 169.

"Les naturalistes out generalemente cru que les animaux microscopiques etaient engendres par des oeufs transportes dans l'air, ou deposes dans des eaux dormantes par des insectes volans."--_Ibid._ p. 176.]

But the skill of the microscope makers of the eighteenth century soon reached its limit. A microscope magnifying 400 diameters was a _chef d'oeuvre_ of the opticians of that day; and, at the same time, by no means trustworthy. But a magnifying power of 400 diameters, even when definition reaches the exquisite perfection of our modern achromatic lenses, hardly suffices for the mere discernment of the smallest forms of life. A speck, only 1/25th of an inch in diameter, has, at ten inches from the eye, the same apparent size as an object 1/10000th of an inch in diameter, when magnified 400 times; but forms of living matter abound, the diameter of which is not more than 1/40000th of an inch. A filtered infusion of hay, allowed to stand for two days, will swarm with living things among which, any which reaches the diameter of a human red blood- corpuscle, or about 1/3200th of an inch, is a giant. It is only by bearing these facts in mind, that we can deal fairly with the remarkable statements and speculations put forward by Buffon and Needham in the middle of the eighteenth century.

When a portion of any animal or vegetable body is infused in water, it gradually softens and disintegrates; and, as it does so, the water is found to swarm with minute active creatures, the so-called Infusorial Animalcules, none of which can be seen, except by the aid of the microscope; while a large proportion belong to the category of smallest things of which I have spoken, and which must have looked like mere dots and lines under the ordinary microscopes of the eighteenth century.

Led by various theoretical considerations which I cannot now discuss, but which looked promising enough in the lights of their time, Buffon and Needham doubted the applicability of Redi's hypothesis to the infusorial animalcules, and Needham very properly endeavoured to put the question to an experimental test. He said to himself, If these infusorial animalcules come from germs, their germs must exist either in the substance infused, or in the water with which the infusion is made, or in the superjacent air. Now the vitality of all germs is destroyed by heat. Therefore, if I boil the infusion, cork it up carefully, cementing the cork over with mastic, and then heat the whole vessel by heaping hot ashes over it, I must needs kill whatever germs are present. Consequently, if Redi's hypothesis hold good, when the infusion is taken away and allowed to cool, no animalcules ought to be developed in it; whereas, if the animalcules are not dependent on pre-existing germs, but are generated from the infused substance, they ought, by and by, to make their appearance. Needham found that, under the circumstances in which he made his experiments, animalcules always did arise in the infusions, when a sufficient time had elapsed to allow for their development.

In much of his work Needham was associated with Buffon, and the results of their experiments fitted in admirably with the great French naturalist's hypothesis of "organic molecules," according to which, life is the indefeasible property of certain indestructible molecules of matter, which exist in all living things, and have inherent activities by which they are distinguished from not living matter. Each individual living organism is formed by their temporary combination. They stand to it in the relation of the particles of water to a cascade, or a whirlpool; or to a mould, into which the water is poured. The form of the organism is thus determined by the reaction between external conditions and the inherent activities of the organic molecules of which it is composed; and, as the stoppage of a whirlpool destroys nothing but a form, and leaves the molecules of the water, with all their inherent activities intact, so what we call the death and putrefaction of an animal, or of a plant, is merely the breaking up of the form, or manner of association, of its constituent organic molecules, which are then set free as infusorial animalcules.

It will be perceived that this doctrine is by no means identical with _Abiogenesis_, with which it is often confounded. On this hypothesis, a piece of beef, or a handful of hay, is dead only in a limited sense. The beef is dead ox, and the hay is dead grass; but the "organic molecules" of the beef or the hay are not dead, but are ready to manifest their vitality as soon as the bovine or herbaceous shrouds in which they are imprisoned are rent by the macerating action of water. The hypothesis therefore must be classified under Xenogenesis, rather than under Abiogenesis. Such as it was, I think it will appear, to those who will be just enough to remember that it was propounded before the birth of modern chemistry, and of the modern optical arts, to be a most ingenious and suggestive speculation.

But the great tragedy of Science--the slaying of a beautiful hypothesis by an ugly fact--which is so constantly being enacted under the eyes of philosophers, was played, almost immediately, for the benefit of Buffon and Needham.

Once more, an Italian, the Abbe Spallanzani, a worthy successor and representative of Redi in his acuteness, his ingenuity, and his learning, subjected the experiments and the conclusions of Needham to a searching criticism. It might be true that Needham's experiments yielded results such as he had described, but did they bear out his arguments? Was it not possible, in the first place, he had not completely excluded the air by his corks and mastic? And was it not possible, in the second place, that he had not sufficiently heated his infusions and the superjacent air? Spallanzani joined issue with the English naturalist on both these pleas, and he showed that if, in the first place, the glass vessels in which the infusions were contained were hermetically sealed by fusing their necks, and if, in the second place, they were exposed to the temperature of boiling water for three-quarters of an hour,[8] no animalcules ever made their appearance within them. It must be admitted that the experiments and arguments of Spallanzani furnish a complete and a crushing reply to those of Needham. But we all too often forget that it is one thing to refute a proposition, and another to prove the truth of a doctrine which, implicitly or explicitly, contradicts that proposition; and the advance of science soon showed that though Needham might be quite wrong, it did not follow that Spallanzani was quite right.

[Footnote 8: See Spallanzani, _Opere_, vi. pp. 42 and 51.]

Modern Chemistry, the birth of the latter half of the eighteenth century, grew apace, and soon found herself face to face with the great problems which biology had vainly tried to attack without her help. The discovery of oxygen led to the laying of the foundations of a scientific theory of respiration, and to an examination of the marvellous interactions of organic substances with oxygen. The presence of free oxygen appeared to be one of the conditions of the existence of life, and of those singular changes in organic matters which are known as fermentation and putrefaction. The question of the generation of the infusory animalcules thus passed into a new phase. For what might not have happened to the organic matter of the infusions, or to the oxygen of the air, in Spallanzani's experiments? What security was there that the development of life which ought to have taken place had not been checked or prevented by these changes?

The battle had to be fought again. It was needful to repeat the experiments under conditions which would make sure that neither the oxygen of the air, nor the composition of the organic matter, was altered in such a manner as to interfere with the existence of life.

Schulze and Schwann took up the question from this point of view in 1836 and 1837. The passage of air through red-hot glass tubes, or through strong sulphuric acid, does not alter the proportion of its oxygen, while it must needs arrest, or destroy, any organic matter which may be contained in the air. These experimenters, therefore, contrived arrangements by which the only air which should come into contact with a boiled infusion should be such as had either passed through red-hot tubes or through strong sulphuric acid. The result which they obtained was that an infusion so treated developed no living things, while, if the same infusion was afterwards exposed to the air, such things appeared rapidly and abundantly. The accuracy of these experiments has been alternately denied and affirmed. Supposing then, to be accepted, however, all that they really proved was that the treatment to which the air was subjected destroyed _something_ that was essential to the development of life in the infusion. This "something" might be gaseous, fluid, or solid; that it consisted of germs remained only an hypothesis of greater or less probability.

Contemporaneously with these investigations a remarkable discovery was made by Cagniard de la Tour. He found that common yeast is composed of a vast accumulation of minute plants. The fermentation of must, or of wort, in the fabrication of wine and of beer, is always accompanied by the rapid growth and multiplication of these _Toruloe_. Thus, fermentation, in so far as it was accompanied by the development of microscopical organisms in enormous numbers, became assimilated to the decomposition of an infusion of ordinary animal or vegetable matter; and it was an obvious suggestion that the organisms were, in some way or other, the causes both of fermentation and of putrefaction. The chemists, with Berzelius and Liebig at their head, at first laughed this idea to scorn; but in 1843, a man then very young, who has since performed the unexampled feat of attaining to high eminence alike in Mathematics, Physics, and Physiology-- I speak of the illustrious Helmholtz--reduced the matter to the test of experiment by a method alike elegant and conclusive. Helmholtz separated a putrefying or a fermenting liquid from one which was simply putrescible or fermentable by a membrane which allowed the fluids to pass through and become intermixed, but stopped the passage of solids. The result was, that while the putrescible or the fermentable liquids became impregnated with the results of the putrescence or fermentation which was going on on the other side of the membrane, they neither putrefied (in the ordinary way) nor fermented; nor were any of the organisms which abounded in the fermenting or putrefying liquid generated in them. Therefore the cause of the development of these organisms must lie in something which cannot pass through membranes; and as Helmholtz's investigations were long antecedent to Graham's researches upon colloids, his natural conclusion was that the agent thus intercepted must be a solid material. In point of fact, Helmholtz's experiments narrowed the issue to this: that which excites fermentation and putrefaction, and at the same time gives rise to living forms in a fermentable or putrescible fluid, is not a gas and is not a diffusible fluid; therefore it is either a colloid, or it is matter divided into very minute solid particles.

The researches of Schroeder and Dusch in 1854, and of Schroeder alone, in 1859, cleared up this point by experiments which are simply refinements upon those of Redi. A lump of cotton-wool is, physically speaking, a pile of many thicknesses of a very fine gauze, the fineness of the meshes of which depends upon the closeness of the compression of the wool. Now, Schroeder and Dusch found, that, in the case of all the putrefiable materials which they used (except milk and yolk of egg), an infusion boiled, and then allowed to come into contact with no air but such as had been filtered through cotton-wool, neither putrefied, nor fermented, nor developed living forms. It is hard to imagine what the fine sieve formed by the cotton-wool could have stopped except minute solid particles. Still the evidence was incomplete until it had been positively shown, first, that ordinary air does contain such particles; and, secondly, that filtration through cotton-wool arrests these particles and allows only physically pure air to pass. This demonstration has been furnished within the last year by the remarkable experiments of Professor Tyndall. It has been a common objection of Abiogenists that, if the doctrine of Biogeny is true, the air must be thick with germs; and they regard this as the height of absurdity. But nature occasionally is exceedingly unreasonable, and Professor Tyndall has proved that this particular absurdity may nevertheless be a reality. He has demonstrated that ordinary air is no better than a sort of stirabout of excessively minute solid particles; that these particles are almost wholly destructible by heat; and that they are strained off, and the air rendered optically pure, by its being passed through cotton-wool.

It remains yet in the order of logic, though not of history, to show that among these solid destructible particles, there really do exist germs capable of giving rise to the development of living forms in suitable menstrua. This piece of work was done by M. Pasteur in those beautiful researches which will ever render his name famous; and which, in spite of all attacks upon them, appear to me now, as they did seven years ago,[9] to be models of accurate experimentation and logical reasoning. He strained air through cotton-wool, and found, as Schroeder and Dusch had done, that it contained nothing competent to give rise to the development of life in fluids highly fitted for that purpose. But the important further links in the chain of evidence added by Pasteur are three. In the first place he subjected to microscopic examination the cotton-wool which had served as strainer, and found that sundry bodies clearly recognisable as germs, were among the solid particles strained off. Secondly, he proved that these germs were competent to give rise to living forms by simply sowing them in a solution fitted for their development. And, thirdly, he showed that the incapacity of air strained through cotton- wool to give rise to life, was not due to any occult change effected in the constituents of the air by the wool, by proving that the cotton-wool might be dispensed with altogether, and perfectly free access left between the exterior air and that in the experimental flask. If the neck of the flask is drawn out into a tube and bent downwards; and if, after the contained fluid has been carefully boiled, the tube is heated sufficiently to destroy any germs which may be present in the air which enters as the fluid cools, the apparatus may be left to itself for any time and no life will appear in the fluid. The reason is plain. Although there is free communication between the atmosphere laden with germs and the germless air in the flask, contact between the two takes place only in the tube; and as the germs cannot fall upwards, and there are no currents, they never reach the interior of the flask. But if the tube be broken short off where it proceeds from the flask, and free access be thus given to germs falling vertically out of the air, the fluid, which has remained clear and desert for months, becomes, in a few days, turbid and full of life.

[Footnote 9: _Lectures to Working Men on the Causes of the Phenomena of Organic Nature_, 1863. (See Vol. II. of these Essays.)]

These experiments have been repeated over and over again by independent observers with entire success; and there is one very simple mode of seeing the facts for one's self, which I may as well describe.

Prepare a solution (much used by M. Pasteur, and often called "Pasteur's solution") composed of water with tartrate of ammonia, sugar, and yeast- ash dissolved therein.[10] Divide it into three portions in as many flasks; boil all three for a quarter of an hour; and, while the steam is passing out, stop the neck of one with a large plug of cotton-wool, so that this also may be thoroughly steamed. Now set the flasks aside to cool, and, when their contents are cold, add to one of the open ones a drop of filtered infusion of hay which has stood for twenty-four hours, and is consequently hill of the active and excessively minute organisms known as _Bacteria_. In a couple of days of ordinary warm weather the contents of this flask will be milky from the enormous multiplication of _Bacteria_. The other flask, open and exposed to the air, will, sooner or later, become milky with _Bacteria_, and patches of mould may appear in it; while the liquid in the flask, the neck of which is plugged with cotton-wool, will remain clear for an indefinite time. I have sought in vain for any explanation of these facts, except the obvious one, that the air contains germs competent to give rise to _Bacteria_, such as those with which the first solution has been knowingly and purposely inoculated, and to the mould-_Fungi_. And I have not yet been able to meet with any advocate of Abiogenesis who seriously maintains that the atoms of sugar, tartrate of ammonia, yeast-ash, and water, under no influence but that of free access of air and the ordinary temperature, re-arrange themselves and give rise to the protoplasm of _Bacterium_. But the alternative is to admit that these _Bacteria_ arise from germs in the air; and if they are thus propagated, the burden of proof that other like forms are generated in a different manner, must rest with the assertor of that proposition.

[Footnote 10: Infusion of hay treated in the same way yields similar results; but as it contains organic matter, the argument which follows cannot be based upon it.]

To sum up the effect of this long chain of evidence:--

It is demonstrable that a fluid eminently fit for the development of the lowest forms of life, but which contains neither germs, nor any protein compound, gives rise to living things in great abundance if it is exposed to ordinary air; while no such development takes place, if the air with which it is in contact is mechanically freed from the solid particles which ordinarily float in it, and which may be made visible by appropriate means.

It is demonstrable that the great majority of these particles are destructible by heat, and that some of them are germs, or living particles, capable of giving rise to the same forms of life as those which appear when the fluid is exposed to unpurified air.

It is demonstrable that inoculation of the experimental fluid with a drop of liquid known to contain living particles gives rise to the same phenomena as exposure to unpurified air.

And it is further certain that these living particles are so minute that the assumption of their suspension in ordinary air presents not the slightest difficulty. On the contrary, considering their lightness and the wide diffusion of the organisms which produce them, it is impossible to conceive that they should not be suspended in the atmosphere in myriads.

Thus the evidence, direct and indirect, in favour of _Biogenesis_ for all known forms of life must, I think, be admitted to be of great weight.

On the other side, the sole assertions worthy of attention are that hermetically sealed fluids, which have been exposed to great and long- continued heat, have sometimes exhibited living forms of low organisation when they have been opened.

The first reply that suggests itself is the probability that there must be some error about these experiments, because they are performed on an enormous scale every day with quite contrary results. Meat, fruits, vegetables, the very materials of the most fermentable and putrescible infusions, are preserved to the extent, I suppose I may say, of thousands of tons every year, by a method which is a mere application of Spallanzani's experiment. The matters to be preserved are well boiled in a tin case provided with a small hole, and this hole is soldered up when all the air in the case has been replaced by steam. By this method they may be kept for years without putrefying, fermenting, or getting mouldy. Now this is not because oxygen is excluded, inasmuch as it is now proved that free oxygen is not necessary for either fermentation or putrefaction. It is not because the tins are exhausted of air, for _Vibriones_ and _Bacteria_ live, as Pasteur has shown, without air or free oxygen. It is not because the boiled meats or vegetables are not putrescible or fermentable, as those who have had the misfortune to be in a ship supplied with unskilfully closed tins well know. What is it, therefore, but the exclusion of germs? I think that Abiogenists are bound to answer this question before they ask us to consider new experiments of precisely the same order.

And in the next place, if the results of the experiments I refer to are really trustworthy, it by no means follows that Abiogenesis has taken place. The resistance of living matter to heat is known to vary within considerable limits, and to depend, to some extent, upon the chemical and physical qualities of the surrounding medium. But if, in the present state of science, the alternative is offered us,--either germs can stand a greater heat than has been supposed, or the molecules of dead matter, for no valid or intelligible reason that is assigned, are able to re- arrange themselves into living bodies, exactly such as can be demonstrated to be frequently produced in another way,--I cannot understand how choice can be, even for a moment, doubtful.

But though I cannot express this conviction of mine too strongly, I must carefully guard myself against the supposition that I intend to suggest that no such thing as Abiogenesis ever has taken place in the past, or ever will take place in the future. With organic chemistry, molecular physics, and physiology yet in their infancy, and every day making prodigious strides, I think it would be the height of presumption for any man to say that the conditions under which matter assumes the properties we call "vital" may not, some day, be artificially brought together. All I feel justified in affirming is, that I see no reason for believing that the feat has been performed yet.

And looking back through the prodigious vista of the past, I find no record of the commencement of life, and therefore I am devoid of any means of forming a definite conclusion as to the conditions of its appearance. Belief, in the scientific sense of the word, is a serious matter, and needs strong foundations. To say, therefore, in the admitted absence of evidence, that I have any belief as to the mode in which the existing forms of life have originated, would be using words in a wrong sense. But expectation is permissible where belief is not; and if it were given me to look beyond the abyss of geologically recorded time to the still more remote period when the earth was passing through physical and chemical conditions, which it can no more see again than a man can recall his infancy, I should expect to be a witness of the evolution of living protoplasm from not living matter. I should expect to see it appear under forms of great simplicity, endowed, like existing fungi, with the power of determining the formation of new protoplasm from such matters as ammonium carbonates, oxalates and tartrates, alkaline and earthy phosphates, and water, without the aid of light. That is the expectation to which analogical reasoning leads me; but I beg you once more to recollect that I have no right to call my opinion anything but an act of philosophical faith.

So much for the history of the progress of Redi's great doctrine of Biogenesis, which appears to me, with the limitations I have expressed, to be victorious along the whole line at the present day.

As regards the second problem offered to us by Redi, whether Xenogenesis obtains, side by side with Homogenesis,--whether, that is, there exist not only the ordinary living things, giving rise to offspring which run through the same cycle as themselves, but also others, producing offspring which are of a totally different character from themselves,-- the researches of two centuries have led to a different result. That the grubs found in galls are no product of the plants on which the galls grow, but are the result of the introduction of the eggs of insects into the substance of these plants, was made out by Vallisnieri, Reaumur, and others, before the end of the first half of the eighteenth century. The tapeworms, bladderworms, and flukes continued to be a stronghold of the advocates of Xenogenesis for a much longer period. Indeed, it is only within the last thirty years that the splendid patience of Von Siebold, Van Beneden, Leuckart, Kuechenmeister, and other helminthologists, has succeeded in tracing every such parasite, often through the strangest wanderings and metamorphoses, to an egg derived from a parent, actually or potentially like itself; and the tendency of inquiries elsewhere has all been in the same direction. A plant may throw off bulbs, but these, sooner or later, give rise to seeds or spores, which develop into the original form. A polype may give rise to Medusae, or a pluteus to an Echinoderm, but the Medusa and the Echinoderm give rise to eggs which produce polypes or glutei, and they are therefore only stages in the cycle of life of the species.

But if we turn to pathology, it offers us some remarkable approximations to true Xenogenesis.

As I have already mentioned, it has been known since the time of Vallisnieri and of Reaumur, that galls in plants, and tumours in cattle, are caused by insects, which lay their eggs in those parts of the animal or vegetable frame of which these morbid structures are outgrowths. Again, it is a matter of familiar experience to everybody that mere pressure on the skin will give rise to a corn. Now the gall, the tumour, and the corn are parts of the living body, which have become, to a certain degree, independent and distinct organisms. Under the influence of certain external conditions, elements of the body, which should have developed in due subordination to its general plan, set up for themselves and apply the nourishment which they receive to their own purposes.

From such innocent productions as corns and warts, there are all gradations to the serious tumours which, by their mere size and the mechanical obstruction they cause, destroy the organism out of which they are developed; while, finally, in those terrible structures known as cancers, the abnormal growth has acquired powers of reproduction and multiplication, and is only morphologically distinguishable from the parasitic worm, the life of which is neither more nor less closely bound up with that of the infested organism.

If there were a kind of diseased structure, the histological elements of which were capable of maintaining a separate and independent existence out of the body, it seems to me that the shadowy boundary between morbid growth and Xenogenesis would be effaced. And I am inclined to think that the progress of discovery has almost brought us to this point already. I have been favoured by Mr. Simon with an early copy of the last published of the valuable "Reports on the Public Health," which, in his capacity of their medical officer, he annually presents to the Lords of the Privy Council. The appendix to this report contains an introductory essay "On the Intimate Pathology of Contagion," by Dr. Burdon-Sanderson, which is one of the clearest, most comprehensive, and well-reasoned discussions of a great question which has come under my notice for a long time. I refer you to it for details and for the authorities for the statements I am about to make.

You are familiar with what happens in vaccination. A minute cut is made in the skin, and an infinitesimal quantity of vaccine matter is inserted into the wound. Within a certain time a vesicle appears in the place of the wound, and the fluid which distends this vesicle is vaccine matter, in quantity a hundred or a thousandfold that which was originally inserted. Now what has taken place in the course of this operation? Has the vaccine matter, by its irritative property, produced a mere blister, the fluid of which has the same irritative property? Or does the vaccine matter contain living particles, which have grown and multiplied where they have been planted? The observations of M. Chauveau, extended and confirmed by Dr. Sanderson himself, appear to leave no doubt upon this head. Experiments, similar in principle to those of Helmholtz on fermentation and putrefaction, have proved that the active element in the vaccine lymph is non-diffusible, and consists of minute particles not exceeding 1/20000th of an inch in diameter, which are made visible in the lymph by the microscope. Similar experiments have proved that two of the most destructive of epizootic diseases, sheep-pox and glanders, are also dependent for their existence and their propagation upon extremely small living solid particles, to which the title of _microzymes_ is applied. An animal suffering under either of these terrible diseases is a source of infection and contagion to others, for precisely the same reason as a tub of fermenting beer is capable of propagating its fermentation by "infection," or "contagion," to fresh wort. In both cases it is the solid living particles which are efficient; the liquid in which they float, and at the expense of which they live, being altogether passive.

Now arises the question, are these microzymes the results of _Homogenesis_, or of _Xenogenesis?_ are they capable, like the _Toruloe_ of yeast, of arising only by the development of pre-existing germs? or may they be, like the constituents of a nut-gall, the results of a modification and individualisation of the tissues of the body in which they are found, resulting from the operation of certain conditions? Are they parasites in the zoological sense, or are they merely what Virchow has called "heterologous growths"? It is obvious that this question has the most profound importance, whether we look at it from a practical or from a theoretical point of view. A parasite may be stamped out by destroying its germs, but a pathological product can only be annihilated by removing the conditions which give rise to it.

It appears to me that this great problem will have to be solved for each zymotic disease separately, for analogy cuts two ways. I have dwelt upon the analogy of pathological modification, which is in favour of the xenogenetic origin of microzymes; but I must now speak of the equally strong analogies in favour of the origin of such pestiferous particles by the ordinary process of the generation of like from like.

It is, at present, a well-established fact that certain diseases, both of plants and of animals, which have all the characters of contagious and infectious epidemics, are caused by minute organisms. The smut of wheat is a well-known instance of such a disease, and it cannot be doubted that the grape-disease and the potato-disease fall under the same category. Among animals, insects are wonderfully liable to the ravages of contagious and infectious diseases caused by microscopic _Fungi_.

In autumn, it is not uncommon to see flies motionless upon a window-pane, with a sort of magic circle, in white, drawn round them. On microscopic examination, the magic circle is found to consist of innumerable spores, which have been thrown off in all directions by a minute fungus called _Empusa muscoe_, the spore-forming filaments of which stand out like a pile of velvet from the body of the fly. These spore-forming filaments are connected with others which fill the interior of the fly's body like so much fine wool, having eaten away and destroyed the creature's viscera. This is the full-grown condition of the _Empusa_. If traced back to its earliest stages, in flies which are still active, and to all appearance healthy, it is found to exist in the form of minute corpuscles which float in the blood of the fly. These multiply and lengthen into filaments, at the expense of the fly's substance; and when they have at last killed the patient, they grow out of its body and give off spores. Healthy flies shut up with diseased ones catch this mortal disease, and perish like the others. A most competent observer, M. Cohn, who studied the development of the _Empusa_ very carefully, was utterly unable to discover in what manner the smallest germs of the _Empusa_ got into the fly. The spores could not be made to give rise to such germs by cultivation; nor were such germs discoverable in the air, or in the food of the fly. It looked exceedingly like a case of Abiogenesis, or, at any rate, of Xenogenesis; and it is only quite recently that the real course of events has been made out. It has been ascertained, that when one of the spores falls upon the body of a fly, it begins to germinate, and sends out a process which bores its way through the fly's skin; this, having reached the interior cavities of its body, gives off the minute floating corpuscles which are the earliest stage of the _Empusa_. The disease is "contagious," because a healthy fly coming in contact with a diseased one, from which the spore-bearing filaments protrude, is pretty sure to carry off a spore or two. It is "infectious" because the spores become scattered about all sorts of matter in the neighbourhood of the slain flies.

The silkworm has long been known to be subject to a very fatal and infectious disease called the _Muscardine_. Audouin transmitted it by inoculation. This disease is entirely due to the development of a fungus, _Botrytis Bassiana_, in the body of the caterpillar; and its contagiousness and infectiousness are accounted for in the same way as those of the fly-disease. But, of late years, a still more serious epizootic has appeared among the silkworms; and I may mention a few facts which will give you some conception of the gravity of the injury which it has inflicted on France alone.

The production of silk has been for centuries an important branch of industry in Southern France, and in the year 1853 it had attained such a magnitude that the annual produce of the French sericulture was estimated to amount to a tenth of that of the whole world, and represented a money- value of 117,000,000 francs, or nearly five millions sterling. What may be the sum which would represent the money-value of all the industries connected with the working up of the raw silk thus produced, is more than I can pretend to estimate. Suffice it to say, that the city of Lyons is built upon French silk as much as Manchester was upon American cotton before the civil war.

Silkworms are liable to many diseases; and, even before 1853, a peculiar epizootic, frequently accompanied by the appearance of dark spots upon the skin (whence the name of "Pebrine" which it has received), had been noted for its mortality. But in the years following 1853 this malady broke out with such extreme violence, that, in 1858, the silk-crop was reduced to a third of the amount which it had reached in 1853; and, up till within the last year or two, it has never attained half the yield of 1853. This means not only that the great number of people engaged in silk growing are some thirty millions sterling poorer than they might have been; it means not only that high prices have had to be paid for imported silkworm eggs, and that, after investing his money in them, in paying for mulberry-leaves and for attendance, the cultivator has constantly seen his silkworms perish and himself plunged in ruin; but it means that the looms of Lyons have lacked employment, and that, for years, enforced idleness and misery have been the portion of a vast population which, in former days, was industrious and well-to-do.

In 1858 the gravity of the situation caused the French Academy of Sciences to appoint Commissioners, of whom a distinguished naturalist, M. de Quatrefages, was one, to inquire into the nature of this disease, and, if possible, to devise some means of staying the plague. In reading the Report[11] made by M. de Quatrefages in 1859, it is exceedingly interesting to observe that his elaborate study of the Pebrine forced the conviction upon his mind that, in its mode of occurrence and propagation, the disease of the silkworm is, in every respect, comparable to the cholera among mankind. But it differs from the cholera, and so far is a more formidable malady, in being hereditary, and in being, under some circumstances, contagious as well as infectious.

[Footnote 11: _Etudes sur les Maladies actuelles des Vers a Soie_, p. 53.]

The Italian naturalist, Filippi, discovered in the blood of the silkworms affected by this strange disorder a multitude of cylindrical corpuscles, each about 1/6000th of an inch long. These have been carefully studied by Lebert, and named by him _Panhistophyton_; for the reason that in subjects in which the disease is strongly developed, the corpuscles swarm in every tissue and organ of the body, and even pass into the undeveloped eggs of the female moth. But are these corpuscles causes, or mere concomitants, of the disease? Some naturalists took one view and some another; and it was not until the French Government, alarmed by the continued ravages of the malady, and the inefficiency of the remedies which had been suggested, despatched M. Pasteur to study it, that the question received its final settlement; at a great sacrifice, not only of the time and peace of mind of that eminent philosopher, but, I regret to have to add, of his health.

But the sacrifice has not been in vain. It is now certain that this devastating, cholera-like, Pebrine, is the effect of the growth and multiplication of the _Panhistophyton_ in the silkworm. It is contagious and infectious, because the corpuscles of the _Panhistophyton_ pass away from the bodies of the diseased caterpillars, directly or indirectly, to the alimentary canal of healthy silkworms in their neighbourhood; it is hereditary because the corpuscles enter into the eggs while they are being formed, and consequently are carried within them when they are laid; and for this reason, also, it presents the very singular peculiarity of being inherited only on the mother's side. There is not a single one of all the apparently capricious and unaccountable phenomena presented by the Pebrine, but has received its explanation from the fact that the disease is the result of the presence of the microscopic organism, _Panhistophyton_.

Such being the facts with respect to the Pebrine, what are the indications as to the method of preventing it? It is obvious that this depends upon the way in which the _Panhistophyton_ is generated. If it may be generated by Abiogenesis, or by Xenogenesis, within the silkworm or its moth, the extirpation of the disease must depend upon the prevention of the occurrence of the conditions under which this generation takes place. But if, on the other hand, the _Panhistophyton_ is an independent organism, which is no more generated by the silkworm than the mistletoe is generated by the apple-tree or the oak on which it grows, though it may need the silkworm for its development in the same way as the mistletoe needs the tree, then the indications are totally different. The sole thing to be done is to get rid of and keep away the germs of the _Panhistophyton_. As might be imagined, from the course of his previous investigations, M. Pasteur was led to believe that the latter was the right theory; and, guided by that theory, he has devised a method of extirpating the disease, which has proved to be completely successful wherever it has been properly carried out.

There can be no reason, then, for doubting that, among insects, contagious and infectious diseases, of great malignity, are caused by minute organisms which are produced from pre-existing germs, or by homogenesis; and there is no reason, that I know of, for believing that what happens in insects may not take place in the highest animals. Indeed, there is already strong evidence that some diseases of an extremely malignant and fatal character to which man is subject, are as much the work of minute organisms as is the Pebrine. I refer for this evidence to the very striking facts adduced by Professor Lister in his various well-known publications on the antiseptic method of treatment. It appears to me impossible to rise from the perusal of those publications without a strong conviction that the lamentable mortality which so frequently dogs the footsteps of the most skilful operator, and those deadly consequences of wounds and injuries which seem to haunt the very walls of great hospitals, and are, even now, destroying more men than die of bullet or bayonet, are due to the importation of minute organisms into wounds, and their increase and multiplication; and that the surgeon who saves most lives will be he who best works out the practical consequences of the hypothesis of Redi.

I commenced this Address by asking you to follow me in an attempt to trace the path which has been followed by a scientific idea, in its long and slow progress from the position of a probable hypothesis to that of an established law of nature. Our survey has not taken us into very attractive regions; it has lain, chiefly, in a land flowing with the abominable, and peopled with mere grubs and mouldiness. And it may be imagined with what smiles and shrugs, practical and serious contemporaries of Redi and of Spallanzani may have commented on the waste of their high abilities in toiling at the solution of problems which, though curious enough in themselves, could be of no conceivable utility to mankind.

Nevertheless, you will have observed that before we had travelled very far upon our road, there appeared, on the right hand and on the left, fields laden with a harvest of golden grain, immediately convertible into those things which the most solidly practical men will admit to have value--viz., money and life.

The direct loss to France caused by the Pebrine in seventeen years cannot be estimated at less than fifty millions sterling; and if we add to this what Redi's idea, in Pasteur's hands, has done for the wine-grower and for the vinegar-maker, and try to capitalise its value, we shall find that it will go a long way towards repairing the money losses caused by the frightful and calamitous war of this autumn. And as to the equivalent of Redi's thought in life, how can we over-estimate the value of that knowledge of the nature of epidemic and epizootic diseases, and consequently of the means of checking, or eradicating them, the dawn of which has assuredly commenced?

Looking back no further than ten years, it is possible to select three (1863, 1864, and 1869) in which the total number of deaths from scarlet- fever alone amounted to ninety thousand. That is the return of killed, the maimed and disabled being left out of sight. Why, it is to be hoped that the list of killed in the present bloodiest of all wars will not amount to more than this! But the facts which I have placed before you must leave the least sanguine without a doubt that the nature and the causes of this scourge will, one day, be as well understood as those of the Pebrine are now; and that the long-suffered massacre of our innocents will come to an end.

And thus mankind will have one more admonition that "the people perish for lack of knowledge"; and that the alleviation of the miseries, and the promotion of the welfare, of men must be sought, by those who will not lose their pains, in that diligent, patient, loving study of all the multitudinous aspects of Nature, the results of which constitute exact knowledge, or Science. It is the justification and the glory of this great meeting that it is gathered together for no other object than the advancement of the moiety of science which deals with those phenomena of nature which we call physical. May its endeavours be crowned with a full measure of success!

[The end]
Thomas Henry Huxley's essay: Biogenesis And Abiogenesis

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