Discourses: Biological and Geological — Biogenesis and Abiogenesis Pt 1

BIOGENESIS AND ABIOGENESIS

(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 aequivocâ (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 tunicâ, aliquâaut 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 aëriform 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 più savio, là 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 più prodotto da se medesima nè erba nè albero, nè animale alcuno perfetto o imperfetto che ei se fosse; e che tutto quello, che ne' tempi trapassati è 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, nè 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 ciò, 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, perchè venendo i bachi per dí fuora, e cercando l' alimento, col rodere ci aprono la strada, ed arrivano alla più interna midolla de' frutti e de' legni.

L'altra maniera si è, che io per me stimerei, che non fosse gran fatto disdicevole il credere, che quell' anima o quella virtù, la quale genera i fiori ed i frutti nelle piante viventi, sia quella stessa che generi ancora i bachi di esse piante.

E chi sà, 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 parrà 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; imperciocchè in quello gallozzole, e particolarmente nelle più 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 è 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 è finita di maturare e che è 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, perchè arrivando la mosca nel tempo della primavera, e facendo una piccolissima fessura ne' rami più 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, Réaurnur, 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 à établir, comme une vérité certaine, que toute plante vient do sa sémence spécifique, tout animal d'un oeuf ou de quelque chose d'analogue préexistant dans la plante, ou dans l'animal de même espèce qui l'a produit."

--_Nouvelles Observations_, p. 169. "Les naturalistes out généralemente cru que les animaux microscopiques étaient engendrés par des oeufs transportés dans l'air, ou déposés 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 Abbé 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.