“The most fertile men of science have made blunders, and their consciousness of such slips has been retribution enough; it is only their more sterile critics who delight to dwell too often and too long on such mistakes.” Biometrika, 1901.

Introductory.

On the rediscovery and confirmation of Mendel’s Law by de Vries, Correns, and Tschermak two years ago, it became clear to many naturalists, as it certainly is to me, that we had found a principle which is destined to play a part in the Study of Evolution comparable only with the achievement of Darwin—that after the weary halt of forty years we have at last begun to march.

If we look back on the post-Darwinian period we recognize one notable effort to advance. This effort—fruitful as it proved, memorable as it must ever be—was that made by Galton when he enuntiated his Law of Ancestral Heredity, subsequently modified and restated by Karl Pearson. Formulated after long and laborious inquiry, this principle beyond question gives us an expression including and denoting many phenomena in which previously no regularity had been detected. But to practical naturalists it was evident from the first that there are great groups of facts which could not on any interpretation be brought within the scope of Galton’s Law, and that by no emendation could that Law be extended to reach them. The existence of these phenomena pointed to a different physiological conception of heredity. Now it is precisely this conception that Mendel’s Law enables us to form. Whether the Mendelian principle can be extended so as to include some apparently Galtonian cases is another question, respecting which we have as yet no facts to guide us, but we have certainly no warrant for declaring such an extension to be impossible.

Whatever answer the future may give to that question, it is clear from this moment that every case which obeys the Mendelian principle is removed finally and irretrievably from the operations of the Law of Ancestral Heredity.

At this juncture Professor Weldon intervenes as a professed exponent of Mendel’s work. It is not perhaps to a devoted partisan of the Law of Ancestral Heredity that we should look for the most appreciative exposition of Mendel, but some bare measure of care and accuracy in representation is demanded no less in justice to fine work, than by the gravity of the issue.

Professor Weldon’s article appears in the current number of Biometrika, Vol. I. Pt. II. which reached me on Saturday, Feb. 8. The paper opens with what purports to be a restatement of Mendel’s experiments and results. In this “restatement” a large part of Mendel’s experiments—perhaps the most significant—are not referred to at all. The perfect simplicity and precision of Mendel’s own account are destroyed; with the result that the reader of Professor Weldon’s paper, unfamiliar with Mendel’s own memoir, can scarcely be blamed if he fail to learn the essence of the discovery. Of Mendel’s conception of the hybrid as a distinct entity with characters proper to itself, apart from inheritance—the most novel thing in the whole paper—Professor Weldon gives no word. Upon this is poured an undigested mass of miscellaneous “facts” and statements from which the reader is asked to conclude, first, that a proposition attributed to Mendel regarding dominance of one character is not of “general”52 application, and finally that “all work based on Mendel’s method” is “vitiated” by a “fundamental mistake,” namely “the neglect of ancestry53.”

To find a parallel for such treatment of a great theme in biology we must go back to those writings of the orthodox which followed the appearance of the “Origin of Species.”

On 17th December 1900 I delivered a Report to the Evolution Committee of the Royal Society on the experiments in Heredity undertaken by Miss E. R. Saunders and myself. This report has been offered to the Society for publication and will I understand shortly appear. In it we have attempted to show the extraordinary significance of Mendel’s principle, to point out what in his results is essential and what subordinate, the ways in which the principle can be extended to apply to a diversity of more complex phenomena—of which some are incautiously cited by Professor Weldon as conflicting facts—and lastly to suggest a few simple terms without which (or some equivalents) the discussion of such phenomena is difficult. Though it is impossible here to give an outline of facts and reasoning there set out at length, I feel that his article needs an immediate reply. Professor Weldon is credited with exceptional familiarity with these topics, and his paper is likely to be accepted as a sufficient statement of the case. Its value will only be known to those who have either worked in these fields themselves or have been at the trouble of thoughtfully studying the original materials.

The nature of Professor Weldon’s article may be most readily indicated if I quote the summary of it issued in a paper of abstracts sent out with Review copies of the Part. This paper was most courteously sent to me by an editor of Biometrika in order to call my attention to the article on Mendel, a subject in which he knew me to be interested. The abstract is as follows.

Such is the judgment a fellow-student passes on this mind

“Voyaging through strange seas of thought alone.”

The only conclusion which most readers could draw from this abstract and indeed from the article it epitomizes, is that Mendel’s discovery so far from being of paramount importance, rests on a basis which Professor Weldon has shown to be insecure, and that an error has come in through disregard of the law of Ancestral Heredity. On examining the paper it is perfectly true that Professor Weldon is careful nowhere directly to question Mendel’s facts or his interpretation of them, for which indeed in some places he even expresses a mild enthusiasm, but there is no mistaking the general purpose of the paper. It must inevitably produce the impression that the importance of the work has been greatly exaggerated and that supporters of current views on Ancestry may reassure themselves. That this is Professor Weldon’s own conclusion in the matter is obvious. After close study of his article it is evident to me that Professor Weldon’s criticism is baseless and for the most part irrelevant, and I am strong in the conviction that the cause which will sustain damage from this debate is not that of Mendel.

I. The Mendelian Principle of Purity of Germ-Cells and the Laws of Heredity Based on Ancestry.

Professor Weldon’s article is entitled “Mendel’s Laws of Alternative Inheritance in Peas.” This title expresses the scope of Mendel’s work and discovery none too precisely and even exposes him to distinct misconception.

To begin with, it says both too little and too much. Mendel did certainly determine Laws of Inheritance in peas—not precisely the laws Professor Weldon has been at the pains of drafting, but of that anon. Having done so, he knew what his discovery was worth. He saw, and rightly, that he had found a principle which must govern a wide area of phenomena. He entitles his paper therefore “Versuche über Pflanzen-Hybriden,” or, Experiments in Plant-Hybridisation.

Nor did Mendel start at first with any particular intention respecting Peas. He tells us himself that he wanted to find the laws of inheritance in hybrids, which he suspected were definite, and that after casting about for a suitable subject, he found one in peas, for the reasons he sets out.

In another respect the question of title is much more important. By the introduction of the word “Alternative” the suggestion is made that the Mendelian principle applies peculiarly to cases of “alternative” inheritance. Mendel himself makes no such limitation in his earlier paper, though perhaps by rather remote implication in the second, to which the reader should have been referred. On the contrary, he wisely abstains from prejudicial consideration of unexplored phenomena.

To understand the significance of the word “alternative” as introduced by Professor Weldon we must go back a little in the history of these studies. In the year 1897 Galton formally announced the Law of Ancestral Heredity referred to in the Introduction, having previously “stated it briefly and with hesitation” in Natural Inheritance, p. 134. In 1898 Professor Pearson published his modification and generalisation of Galton’s Law, introducing a correction of admitted theoretical importance, though it is not in question that the principle thus restated is fundamentally not very different from Galton’s54. It is an essential part of the Galton-Pearson Law of Ancestral Heredity that in calculating the probable structure of each descendant the structure of each several ancestor must be brought to account.

Professor Weldon now tells us that these two papers of Galton and of Professor Pearson have “given us an expression for the effects of blended inheritance which seems likely to prove generally applicable, though the constants of the equations which express the relation between divergence from the mean in one generation, and that in another, may require modification in special cases. Our knowledge of particulate or mosaic inheritance, and of alternative inheritance, is however still rudimentary, and there is so much contradiction between the results obtained by different observers, that the evidence available is difficult to appreciate.”

But Galton stated (p. 401) in 1897 that his statistical law of heredity “appears to be universally applicable to bi-sexual descent.” Pearson in re-formulating the principle in 1898 made no reservation in regard to “alternative” inheritance. On the contrary he writes (p. 393) that “if Mr Galton’s law can be firmly established, it is a complete solution, at any rate to a first approximation, of the whole problem of heredity,” and again (p. 412) that “it is highly probable that it [this law] is the simple descriptive statement which brings into a single focus all the complex lines of hereditary influence. If Darwinian evolution be natural selection combined with heredity, then the single statement which embraces the whole field of heredity must prove almost as epoch-making as the law of gravitation to the astronomer55.”

As I read there comes into my mind that other fine passage where Professor Pearson warns us

No naturalist who had read Galton’s paper and had tried to apply it to the facts he knew could fail to see that here was a definite advance. We could all perceive phenomena that were in accord with it and there was no reasonable doubt that closer study would prove that accord to be close. It was indeed an occasion for enthusiasm, though no one acquainted with the facts of experimental breeding could consider the suggestion of universal application for an instant.

But two years have gone by, and in 1900 Pearson writes57 that the values obtained from the Law of Ancestral Heredity

or, again in other words, that the Law of Ancestral Heredity after the glorious launch in 1898 has been home for a complete refit. The top-hamper is cut down and the vessel altogether more manageable; indeed she looks trimmed for most weathers. Each of the qualifications now introduced wards off whole classes of dangers. Later on (pp. 487–8) Pearson recites a further list of cases regarded as exceptional. “All characters will be inherited at the same rate” might indeed almost be taken to cover the results in Mendelian cases, though the mode by which those results are arrived at is of course wholly different.

Clearly we cannot speak of the Law of Gravitation now. Our Tycho Brahe and our Kepler, with the yet more distant Newton, are appropriately named as yet to come58.

But the truth is that even in 1898 such a comparison was scarcely happy. Not to mention moderns, these high hopes had been finally disposed of by the work of the experimental breeders such as Kölreuter, Knight, Herbert, Gärtner, Wichura, Godron, Naudin, and many more. To have treated as non-existent the work of this group of naturalists, who alone have attempted to solve the problems of heredity and species—Evolution, as we should now say—by the only sound method—experimental breeding—to leave out of consideration almost the whole block of evidence collected in Animals and Plants—Darwin’s finest legacy as I venture to declare—was unfortunate on the part of any exponent of Heredity, and in the writings of a professed naturalist would have been unpardonable. But even as modified in 1900 the Law of Ancestral Heredity is heavily over-sparred, and any experimental breeder could have increased Pearson’s list of unconformable cases by as many again.

But to return to Professor Weldon. He now repeats that the Law of Ancestral Heredity seems likely to prove generally applicable to blended inheritance, but that the case of alternative inheritance is for the present reserved. We should feel more confidence in Professor Weldon’s exposition if he had here reminded us that the special case which fitted Galton’s Law so well that it emboldened him to announce that principle as apparently “universally applicable to bi-sexual descent” was one of alternative inheritance—namely the coat-colour of Basset-hounds. Such a fact is, to say the least, ominous. Pearson, in speaking (1900) of this famous case of Galton’s, says that these phenomena of alternative inheritance must be treated separately (from those of blended inheritance)59, and for them he deduces a proposed “law of reversion,” based of course on ancestry. He writes, “In both cases we may speak of a law of ancestral heredity, but the first predicts the probable character of the individual produced by a given ancestry, while the second tells us the percentages of the total offspring which on the average revert to each ancestral type60.”

With the distinctions between the original Law of Ancestral Heredity, the modified form of the same law, and the Law of Reversion, important as all these considerations are, we are not at present concerned.

For the Mendelian principle of heredity asserts a proposition absolutely at variance with all the laws of ancestral heredity, however formulated. In those cases to which it applies strictly, this principle declares that the cross-breeding of parents need not diminish the purity of their germ-cells or consequently the purity of their offspring. When in such cases individuals bearing opposite characters, A and B, are crossed, the germ-cells of the resulting cross-bred, AB, are each to be bearers either of character A or of character B, not both.

Consequently when the cross-breds breed either together or with the pure forms, individuals will result of the forms AA, AB, BA, BB61. Of these the forms AA and BB, formed by the union of similar germs, are stated to be as pure as if they had had no cross in their pedigree, and henceforth their offspring will be no more likely to depart from the A type or the B type respectively, than those of any other originally pure specimens of these types.

Consequently in such examples it is not the fact that each ancestor must be brought to account as the Galton-Pearson Law asserts, and we are clearly dealing with a physiological phenomenon not contemplated by that Law at all.

Every case therefore which obeys the Mendelian principle is in direct contradiction to the proposition to which Professor Weldon’s school is committed, and it is natural that he should be disposed to consider the Mendelian principle as applying especially to “alternative” inheritance, while the law of Galton and Pearson is to include the phenomenon of blended inheritance. The latter, he tells us, is “the most usual case,” a view which, if supported by evidence, might not be without value.

It is difficult to blame those who on first acquaintance concluded Mendel’s principle can have no strict application save to alternative inheritance. Whatever blame there is in this I share with Professor Weldon and those whom he follows. Mendel’s own cases were almost all alternative; also the fact of dominance is very dazzling at first. But that was two years ago, and when one begins to see clearly again, it does not look so certain that the real essence of Mendel’s discovery, the purity of germ-cells in respect of certain characters, may not apply also to some phenomena of blended inheritance. The analysis of this possibility would take us to too great length, but I commend to those who are more familiar with statistical method, the consideration of this question: whether dominance being absent, indefinite, or suppressed, the phenomena of heritages completely blended in the zygote, may not be produced by gametes presenting Mendelian purity of characters. A brief discussion of this possibility is given in the Introduction, p. 31.

Very careful inquiry would be needed before such a possibility could be negatived. For example, we know that the Laws based on Ancestry can apply to alternative inheritance; witness the case of the Basset-hounds. Here there is no simple Mendelian dominance; but are we sure there is no purity of germ-cells? The new conception goes a long way and it may well reach to such facts as these.

But for the present we will assume that Mendel’s principle applies only to certain phenomena of alternative inheritance, which is as far as our warrant yet runs.

No close student of the recent history of evolutionary thought needs to be told what the attitude of Professor Weldon and his followers has been towards these same disquieting and unwelcome phenomena of alternative inheritance and discontinuity in variation. Holding at first each such fact for suspect, then treating them as rare and negligible occurrences, he and his followers have of late come slowly to accede to the facts of discontinuity a bare and grudging recognition in their scheme of evolution62.

Therefore on the announcement of that discovery which once and for all ratifies and consolidates the conception of discontinuous variation, and goes far to define that of alternative inheritance, giving a finite body to what before was vague and tentative, it is small wonder if Professor Weldon is disposed to criticism rather than to cordiality.

We have now seen what is the essence of Mendel’s discovery based on a series of experiments of unequalled simplicity which Professor Weldon does not venture to dispute.

II. Mendel and the Critic’s Version of him.

The “Law of Dominance.”

I proceed to the question of dominance which Professor Weldon treats as a prime issue, almost to the virtual concealment of the great fact of gametic purity.

Cross-breds in general, AB and BA, named above, may present many appearances. They may all be indistinguishable from A, or from B; some may appear A’s and some B’s; they may be patchworks of both; they may be blends presenting one or many grades between the two; and lastly they may have an appearance special to themselves (being in the latter case, as it often happens, “reversionary”), a possibility which Professor Weldon does not stop to consider, though it is the clue that may unravel many of the facts which mystify him now.

Mendel’s discovery became possible because he worked with regular cases of the first category, in which he was able to recognize that one of each of the pairs of characters he studied did thus prevail and was “dominant” in the cross-bred to the exclusion of the other character. This fact, which is still an accident of particular cases, Professor Weldon, following some of Mendel’s interpreters, dignifies by the name of the “Law of Dominance,” though he omits to warn his reader that Mendel states no “Law of Dominance” whatever. The whole question whether one or other character of the antagonistic pair is dominant though of great importance is logically a subordinate one. It depends on the specific nature of the varieties and individuals used, sometimes probably on the influence of external conditions and on other factors we cannot now discuss. There is as yet no universal law here perceived or declared.

Professor Weldon passes over the proof of the purity of the germ-cells lightly enough, but this proposition of dominance, suspecting its weakness, he puts prominently forward. Briefest equipment will suffice. Facing, as he supposes, some new pretender—some local Theudas—offering the last crazy prophecy,—any argument will do for such an one. An eager gathering in an unfamiliar literature, a scrutiny of samples, and he will prove to us with small difficulty that dominance of yellow over green, and round over wrinkled, is irregular even in peas after all; that in the sharpness of the discontinuity exhibited by the variations of peas there are many grades; that many of these grades co-exist in the same variety; that some varieties may perhaps be normally intermediate. All these propositions are supported by the production of a collection of evidence, the quality of which we shall hereafter consider. “Enough has been said,” he writes (p. 240), “to show the grave discrepancy between the evidence afforded by Mendel’s own experiments and that obtained by other observers, equally competent and trustworthy.”

We are asked to believe that Professor Weldon has thus discovered “a fundamental mistake” vitiating all that work, the importance of which, he elsewhere tells us, he has “no wish to belittle.”

III. The Facts in regard to Dominance of Characters in Peas.

Professor Weldon refers to no experiments of his own and presumably has made none. Had he done so he would have learnt many things about dominance in peas, whether of the yellow cotyledon-colour or of the round form, that might have pointed him to caution.

In the year 1900 Messrs Vilmorin-Andrieux & Co. were kind enough to send to the Cambridge Botanic Garden on my behalf a set of samples of the varieties of Pisum and Phaseolus, an exhibit of which had greatly interested me at the Paris Exhibition of that year. In the past summer I grew a number of these and made some preliminary cross-fertilizations among them (about 80 being available for these deductions) with a view to a future study of certain problems, Mendelian and others. In this work I had the benefit of the assistance of Miss Killby of Newnham College. Her cultivations and crosses were made independently of my own, but our results are almost identical. The experience showed me, what a naturalist would expect and practical men know already, that a great deal turns on the variety used; that some varieties are very sensitive to conditions while others maintain their type sturdily; that in using certain varieties Mendel’s experience as to dominance is regularly fulfilled, while in the case of other varieties irregularities and even some contradictions occur. That the dominance of yellow cotyledon-colour over green, and the dominance of the smooth form over the wrinkled, is a general truth for Pisum sativum appears at once; that it is a universal truth I cannot believe any competent naturalist would imagine, still less assert. Mendel certainly never did. When he speaks of the “law” or “laws” that he has established for Pisum he is referring to his own discovery of the purity of the germ-cells, that of the statistical distribution of characters among them, and the statistical grouping of the different germ-cells in fertilization, and not to the “Law of Dominance” which he never drafted and does not propound.

The issue will be clearer if I here state briefly what, as far as my experience goes, are the facts in regard to the characters cotyledon-colour and seed-shapes in peas. I have not opportunity for more than a passing consideration of the seed-coats of pure forms63; that is a maternal character, a fact I am not sure Professor Weldon fully appreciates. Though that may be incredible, it is evident from many passages that he has not, in quoting authorities, considered the consequences of this circumstance.

The normal characters: colour of cotyledons and seed-coats.

Culinary peas (P. sativum, omitting purple sorts) can primarily be classified on colour into two groups, yellow and green. In the green certain pigmentary matters persist in the ripe seed which disappear or are decomposed in the yellow as the seed ripens. But it may be observed that the “green” class itself is treated as of two divisions, green and blue. In the seedsmen’s lists the classification is made on the external appearance of the seed, without regard to whether the colour is due to the seed-coat, the cotyledons, or both. As a rule perhaps yellow coats contain yellow cotyledons, and green coats green cotyledons, though yellow cotyledons in green coats are common, e.g. Gradus, of which the cotyledons are yellow while the seed-coats are about as often green as yellow (or “white,” as it is called technically). Those called “blue” consist mostly of seeds which have green cotyledons seen through transparent skins, or yellow cotyledons combined with green skins. The skins may be roughly classified into thin and transparent, or thick and generally at some stage pigmented. In numerous varieties the colour of the cotyledon is wholly yellow, or wholly green. Next there are many varieties which are constant in habit and other properties but have seeds belonging to these two colour categories in various proportions. How far these proportions are known to be constant I cannot ascertain.

Of such varieties showing mixture of cotyledon-colours nearly all can be described as dimorphic in colour. For example in Sutton’s Nonpareil Marrowfat the cotyledons are almost always either yellow or green, with some piebalds, and the colours of the seed-coats are scarcely less distinctly dimorphic. In some varieties which exist in both colours intermediates are so common that one cannot assert any regular dimorphism64.

There are some varieties which have cotyledons green and intermediate shading to greenish yellow, like Stratagem quoted by Professor Weldon. Others have yellow and intermediate shading to yellowish green, such as McLean’s Best of all65‍. I am quite disposed to think there may be truly monomorphic varieties with cotyledons permanently of intermediate colour only, but so far I have not seen one66. The variety with greatest irregularity (apart from regular dimorphism) in cotyledon-colour I have seen is a sample of “mange-tout à rames, à grain vert,” but it was a good deal injured by weevils (Bruchus), which always cause irregularity or change of colour.

Lastly in some varieties there are many piebalds or mosaics.

From what has been said it will be evident that the description of a pea in an old book as having been green, blue, white, and so forth, unless the cotyledon-colour is distinguished from seed-coat colour, needs careful consideration before inferences are drawn from it.

Shape.

In regard to shape, if we keep to ordinary shelling peas, the facts are somewhat similar, but as shape is probably more sensitive to conditions than cotyledon-colour (not than seed-coat colour) there are irregularities to be perhaps ascribed to this cause. Broadly, however, there are two main divisions, round and wrinkled. It is unquestioned that between these two types every intermediate occurs. Here again a vast number of varieties can be at once classified into round and wrinkled (the classification commonly used), others are intermediate normally. Here also I suspect some fairly clear sub-divisions might be made in the wrinkled group and in the round group too, but I would not assert this as a fact.

I cannot ascertain from botanists what is the nature of the difference between round and wrinkled peas, though no doubt it will be easily discovered. In maize the round seeds contain much unconverted starch, while in the wrinkled or sugar-maize this seems to be converted in great measure as the seed ripens; with the result that, on drying, the walls collapse. In such seeds we may perhaps suppose that the process of conversion, which in round seeds takes place on germination, is begun earlier, and perhaps the variation essentially consists in the premature appearance of the converting ferment. It would be most rash to suggest that such a process may be operating in the pea, for the phenomenon may have many causes; but however that may be, there is evidently a difference of such a nature that when the water dries out of the seed on ripening, its walls collapse67; and this collapse may occur in varying degrees.

In respect of shape the seeds of a variety otherwise stable are as a rule fairly uniform, the co-existence of both shapes and of intermediates between them in the same variety is not infrequent. As Professor Weldon has said, Telephone is a good example of an extreme case of mixture of both colours and shapes. William I. is another. It may be mentioned that regular dimorphism in respect of shape is not so common as dimorphism in respect of colour. Of great numbers of varieties seen at Messrs Suttons’ I saw none so distinctly dimorphic in shape as William I. which nevertheless contains all grades commonly.

So far I have spoken of the shapes of ordinary English culinary peas. But if we extend our observations to the shapes of large-seeded peas, which occur for the most part among the sugar-peas (mange-touts), of the “grey” peas with coloured flowers, etc., there are fresh complications to be considered.

Professor Weldon does not wholly avoid these (as Mendel did in regard to shape) and we will follow him through his difficulties hereafter. For the present let me say that the classes round and wrinkled are not readily applicable to those other varieties and are not so applied either by Mendel or other practical writers on these subjects. To use the terms indicated in the Introduction, seed-shape depends on more than one pair of allelomorphs—possibly on several.

Stability and Variability.

Generally speaking peas which when seen in bulk are monomorphic in colour and shape, will give fairly true and uniform offspring (but such strict monomorphism is rather exceptional). Instances to the contrary occur, and in my own brief experience I have seen some. In a row of Fill-basket grown from selected seed there were two plants of different habit, seed-shape, etc. Each bore pods with seeds few though large and round. Again Blue Peter (blue and round) and Laxton’s Alpha (blue and wrinkled), grown in my garden and left to nature uncovered, have each given a considerable proportion of seeds with yellow cotyledons, about 20% in the case of Laxton’s Alpha. The distribution of these on the plants I cannot state. The plants bearing them in each case sprang from green-cotyledoned seeds taken from samples containing presumably unselected green seeds only. A part of this exceptional result may be due to crossing, but heterogeneity of conditions68 especially in or after ripening is a more likely cause, hypotheses I hope to investigate next season. Hitherto I had supposed the crossing, if any, to be done by Bruchus or Thrips, but Tschermak also suspects Megachile, the leaf-cutter bee, which abounds in my garden.

Whatever the cause, these irregularities may undoubtedly occur; and if they be proved to be largely independent of crossing and conditions, this will in nowise vitiate the truth of the Mendelian principle. For in that case it may simply be variability. Such true variation, or sporting, in the pea is referred to by many observers. Upon this subject I have received most valuable facts from Mr Arthur Sutton, who has very kindly interested himself in these inquiries. He tells me that several highly bred varieties, selected with every possible care, commonly throw a small but constant proportion of poor and almost vetch-like plants, with short pods and small round seeds, which are hoed out by experienced men each year before ripening. Other high-class varieties always, wherever grown, and when far from other sorts, produce a small percentage of some one or more definite “sports.” Of these peculiar sports he has sent me a collection of twelve, taken from as many standard varieties, each “sport” being represented by eight seeds, which though quite distinct from the type agree with each other in almost all cases.

In two cases, he tells me, these seed-sports sown separately have been found to give plants identical with the standard type and must therefore be regarded as sports in seed characters only; in other cases change of plant-type is associated with the change of seed-type.

In most standard varieties these definite sports are not very common, but in a few they are common enough to require continual removal by selection69.

I hope before long to be able to give statistical details and experiments relating to this extraordinarily interesting subject. As de Vries writes in his fine work Die Mutationstheorie (I. p. 580), “a study of the seed-differences of inconstant, or as they are called, ‘still’ unfixed varieties, is a perfect treasure-house of new discoveries.”

Let us consider briefly the possible significance of these facts in the light of Mendelian teaching. First, then, it is clear that as regards most of such cases the hypothesis is not excluded that these recurring sports may be due to the fortuitous concurrence of certain scarcer hypallelomorphs, which may either have been free in the original parent varieties from which the modern standard forms were raised, or may have been freed in the crossing to which the latter owe their origin (see p. 28). This possibility raises the question whether, if we could make “pure cultures” of the gametes, any variations of this nature would ever occur. This may be regarded as an unwarrantable speculation, but it is not wholly unamenable to the test of experiments.

But variability, in the sense of division of gonads into heterogeneous gametes, may surely be due to causes other than crossing. This we cannot doubt. Cross-fertilization of the zygote producing those gametes is one of the causes of such heterogeneity among them. We cannot suppose it to be the sole cause of this phenomenon.

When Mendel asserts the purity of the germ-cells of cross-breds he cannot be understood to mean that they are more pure than those of the original parental races. These must have varied in the past. The wrinkled seed arose from the round, the green from the yellow (or vice versâ, if preferred), and probably numerous intermediate forms from both.

The variations, or as I provisionally conceive it, that differentiant division among the gametes of which variation (neglecting environment) is the visible expression, has arisen and can arise at one or more points of time, and we have no difficulty in believing it to occur now. In many cases we have clear evidence that it does. Crossing,—dare we call it asymmetrical fertilization?—is one of the causes of the production of heterogeneous gametes—the result of divisions qualitatively differentiant and perhaps asymmetrical70.

There are other causes and we have to find them. Some years ago I wrote that consideration of the causes of variation was in my judgment premature71. Now that through Mendel’s work we are clearing our minds as to the fundamental nature of “gametic” variation, the time is approaching when an investigation of such causes may be not unfruitful.

Of variation as distinct from transmission why does Professor Weldon take no heed? He writes (p. 244):

Now, as I have already pointed out, Mendel made no pretence of universal statement: but had he done so, the conclusion, which Professor Weldon here suggests should follow from such a universal statement, is incorrectly drawn. Mendel is concerned with the laws of transmission of existing characters, not with variation, which he does not discuss.

Nevertheless Professor Weldon has some acquaintance with the general fact of variability in certain peas, which he mentions (p. 236), but the bearing of this fact on the difficulty he enuntiates escapes him.

Results of crossing in regard to seed characters: normal and exceptional.

The conditions being the same, the question of the characters of the cross-bred zygotes which we will call AB’s depends primarily on the specific nature of the varieties which are crossed to produce them. It is unnecessary to point out that if all AB’s are to look alike, both the varieties A and B must be pure—not in the common sense of descended, as far as can be traced, through individuals identical with themselves, but pure in the Mendelian sense, that is to say that each must be at that moment producing only homogeneous gametes bearing the same characters A and B respectively. Purity of pedigree in the breeder’s sense is a distinct matter altogether. The length of time—or if preferred—the number of generations through which a character of a variety has remained pure, alters the probability of its dominance, i.e. its appearance when a gamete bearing it meets another bearing an antagonistic character, no more, so far as we are yet aware, than the length of time a stable element has been isolated alters the properties of the chemical compound which may be prepared from it.

Now when individuals (bearing contrary characters), pure in the sense indicated, are crossed together, the question arises, What will be the appearance of the first cross individuals? Here again, generally speaking, when thoroughly green cotyledons are crossed with thoroughly yellow cotyledons, the first-cross seeds will have yellow cotyledons; when fully round peas are crossed with fully wrinkled the first result will generally speaking be round, often with slight pitting as Mendel has stated. This has been the usual experience of Correns, Tschermak, Mendel, and myself72 and, as we shall see, the amount of clear and substantial evidence to the contrary is still exceedingly small. But as any experienced naturalist would venture to predict, there is no universal rule in the matter. As Professor Weldon himself declares, had there been such a universal rule it would surely have been notorious. He might further have reflected that in Mendel’s day, when hybridisation was not the terra incognita it has since become, the assertion of such universal propositions would have been peculiarly foolish. Mendel does not make it; but Professor Weldon perceiving the inherent improbability of the assertion conceives at once that Mendel must have made it, and if Mendel doesn’t say so in words then he must have implied it. As a matter of fact Mendel never treats dominance as more than an incident in his results, merely using it as a means to an end, and I see no reason to suppose he troubled to consider to what extent the phenomenon is or is not universal—a matter with which he had no concern.

Of course there may be exceptions. As yet we cannot detect the causes which control them, though injury, impurity, accidental crossing, mistakes of various kinds, account for many. Mendel himself says, for instance, that unhealthy or badly grown plants give uncertain results. Nevertheless there seems to be a true residuum of exceptions not to be explained away. I will recite some that I have seen. In my own crosses I have seen green × green give yellow four times. This I incline to attribute to conditions or other disturbance, for the natural pods of these plants gave several yellows. At Messrs Suttons’ I saw second-generation seeds got by allowing a cross of Sutton’s Centenary (gr. wr.) × Eclipse (gr. rd.) to go to seed; the resulting seeds were both green and yellow, wrinkled and round. But in looking at a sample of Eclipse I found a few yellow seeds, say two per cent., which may perhaps be the explanation. Green wrinkled × green round may give all wrinkled, and again wrinkled × wrinkled may give round73. Of this I saw a clear case—supposing no mistake to have occurred—at Messrs Suttons’. Lastly we have the fact that in exceptional cases crossing two forms—apparently pure in the strict sense—may give a mixture in the first generation. There are doubtless examples also of unlikeness between reciprocals, and of this too I have seen one putative case74.

Such facts thus set out for the first cross-bred generation may without doubt be predicated for subsequent generations.

What then is the significance of the facts?

Analysis of exceptions.

Assuming that all these “contradictory” phenomena happened truly as alleged, and were not pathological or due to error—an explanation which seems quite inadequate—there are at least four possible accounts of such diverse results—each valid, without any appeal to ancestry.

1. That dominance may exceptionally fail—or in other words be created on the side which is elsewhere recessive. For this exceptional failure we have to seek exceptional causes. The artificial creation of dominance (in a character usually recessive) has not yet to my knowledge been demonstrated experimentally, but experiments are begun by which such evidence may conceivably be obtained.

2. There may be what is known to practical students of evolution as the false hybridism of Millardet, or in other words, fertilisation with—from unknown causes—transmission of none or of only some of the characters of one pure parent. The applicability of this hypothesis to the colours and shapes of peas is perhaps remote, but we may notice that it is one possible account of those rare cases where two pure forms give a mixed result in the first generation, even assuming the gametes of each pure parent to be truly monomorphic as regards the character they bear. The applicability of this suggestion can of course be tested by study of the subsequent generations, self-fertilised or fertilised by similar forms produced in the same way. In the case of a genuine false-hybrid the lost characters will not reappear in the posterity.

3. The result may not be a case of transmission at all as it is at present conceived, but of the creation on crossing of something new. Our AB’s may have one or more characters peculiar to themselves. We may in fact have made a distinct “mule” or heterozygote form. Where this is the case, there are several subordinate possibilities we need not at present pursue.

4. There may be definite variation (distinct from that proper to the “mule”) consequent on causes we cannot yet surmise (see pp. 125 and 128).

The above possibilities are I believe at the present time the only ones that need to be considered in connexion with these exceptional cases75. They are all of them capable of experimental test and in certain instances we are beginning to expect the conclusion.

The “mule” or heterozygote.

There can be little doubt that in many cases it is to the third category that the phenomena belong. An indication of the applicability of this reasoning will generally be found in the fact that in such “mule” forms the colour or the shape of the seeds will be recognizably peculiar and proper to the specimens themselves, as distinct from their parents, and we may safely anticipate that when those seeds are grown the plants will show some character which is recognizable as novel. The proof that the reasoning may apply can as yet only be got by finding that the forms in question cannot breed true even after successive selections, but constantly break up into the same series of forms76.

This conception of the “mule” form, or “hybrid-character” as Mendel called it, though undeveloped, is perfectly clear in his work. He says that the dominant character may have two significations, it may be either a parental character or a hybrid-character, and it must be differentiated according as it appears in the one capacity or the other. He does not regard the character displayed by the hybrid, whether dominant or other, as a thing inherited from or transmitted by the pure parent at all, but as the peculiar function or property of the hybrid. When this conception has been fully understood and appreciated in all its bearings it will be found to be hardly less fruitful than that of the purity of the germ-cells.

The two parents are two—let us say—substances77 represented by corresponding gametes. These gametes unite to form a new “substance”—the cross-bred zygote. This has its own properties and structure, just as a chemical compound has, and the properties of this new “substance” are not more strictly traceable to, or “inherited” from, those of the two parents than are those of a new chemical compound “inherited” from those of the component elements. If the case be one in which the gametes are pure, the new “substance” is not represented by them, but the compound is again dissociated into its components, each of which is separately represented by gametes.

The character of the cross-bred zygote may be anything. It may be something we have seen before in one or other of the parents, it may be intermediate between the two, or it may be something new. All these possibilities were known to Mendel and he is perfectly aware that his principle is equally applicable to all. The first case is his “dominance.” That he is ready for the second is sufficiently shown by his brief reference to time of flowering considered as a character (p. 65). The hybrids, he says, flower at a time almost exactly intermediate between the flowering times of the parents, and he remarks that the development of the hybrids in this case probably happens in the same way as it does in the case of the other characters78.

That he was thoroughly prepared for the third possibility appears constantly through the paper, notably in the argument based on the Phaseolus hybrids, and in the statement that the hybrid between talls and dwarfs is generally taller than the tall parent, having increased height as its “hybrid-character.”

All this Professor Weldon has missed. In place of it he offers us the sententia that no one can expect to understand these phenomena if he neglect ancestry. This is the idle gloss of the scribe, which, if we erase it not thoroughly, may pass into the text.

Enough has been said to show how greatly Mendel’s conception of heredity was in advance of those which pass current at the present day; I have here attempted the barest outline of the nature of the “hybrid-character,” and I have not sought to indicate the conclusions that we reach when the reasoning so clear in the case of the hybrid is applied to the pure forms and their own characters.

In these considerations we reach the very base on which all conceptions of heredity and variation must henceforth rest, and that it is now possible for us to attempt any such analysis is one of the most far-reaching consequences of Mendel’s principle. Till two years ago no one had made more than random soundings of this abyss.

I have briefly discussed these possibilities to assist the reader in getting an insight into Mendel’s conceptions. But in dealing with Professor Weldon we need not make this excursion; for his objection arising from the absence of uniform regularity in dominance is not in point.

The soundness of Mendel’s work and conclusions would be just as complete if dominance be found to fail often instead of rarely. For it is perfectly certain that varieties can be chosen in such a way that the dominance of one character over its antagonist is so regular a phenomenon that it can be used in the way Mendel indicates. He chose varieties, in fact, in which a known character was regularly dominant and it is because he did so that he made his discovery79. When Professor Weldon speaks of the existence of fluctuation and diversity in regard to dominance as proof of a “grave discrepancy” between Mendel’s facts and those of other observers80, he merely indicates the point at which his own misconceptions began.

From Mendel’s style it may be inferred that if he had meant to state universal dominance in peas he would have done so in unequivocal language. Let me point out further that of the 34 varieties he collected for study, he discarded 12 as not amenable to his purposes81. He tells us he would have nothing to do with characters which were not sharp, but of a “more or less” description. As the 34 varieties are said to have all come true from seed, we may fairly suppose that the reason he discarded twelve was that they were unsuitable for his calculations, having either ill-defined and intermediate characters, or possibly defective and irregular dominance.

IV. Professor Weldon’s collection of “Other Evidence concerning Dominance in Peas.”

A. In regard to cotyledon colour: Preliminary.

I have been at some pains to show how the contradictory results, no doubt sometimes occurring, on which Professor Weldon lays such stress, may be comprehended without any injury to Mendel’s main conclusions. This excursion was made to save trouble with future discoverers of exceptions, though the existence of such facts need scarcely disturb many minds. As regards the dominance of yellow cotyledon-colour over green the whole number of genuine unconformable cases is likely to prove very small indeed, though in regard to the dominance of round shape over wrinkled we may be prepared for more discrepancies. Indeed my own crosses alone are sufficient to show that in using some varieties irregularities are to be expected. Considering also that the shapes of peas depend unquestionably on more than one pair of allelomorphs I fully expect regular blending in some cases.

As however it may be more satisfactory to the reader and to Professor Weldon if I follow him through his “contradictory” evidence I will endeavour to do so. Those who have even a slight practical acquaintance with the phenomena of heredity will sympathize with me in the difficulty I feel in treating this section of his arguments with that gravity he conceives the occasion to demand.

In following the path of the critic it will be necessary for me to trouble the reader with a number of details of a humble order, but the journey will not prove devoid of entertainment.

Now exceptions are always interesting and suggestive things, and sometimes hold a key to great mysteries. Still when a few exceptions are found disobeying rules elsewhere conformed to by large classes of phenomena it is not an unsafe course to consider, with such care as the case permits, whether the exceptions may not be due to exceptional causes, or failing such causes whether there may be any possibility of error. But to Professor Weldon, an exception is an exception—and as such may prove a very serviceable missile; so he gathers them as they were “smooth stones from the brook.”

Before examining the quality of this rather miscellaneous ammunition I would wish to draw the non-botanical reader’s attention to one or two facts of a general nature.

For our present purpose the seed of a pea may be considered as consisting of two parts, the embryo with its cotyledons, enclosed in a seed-coat. It has been known for about a century that this coat or skin is a maternal structure, being part of the mother plant just as much as the pods are, and consequently not belonging to the next generation at all. If then any changes take place in it consequent on fertilisation, they are to be regarded not as in any sense a transmission of character by heredity, but rather as of the nature of an “infection.” If on the other hand it is desired to study the influence of hereditary transmission on seed-coat characters, then the crossed seeds must be sown and the seed-coats of their seeds studied. Such infective changes in maternal tissues have been known from early times, a notable collection of them having been made especially by Darwin; and for these cases Focke suggested the convenient word Xenia. With this familiar fact I would not for a moment suppose Professor Weldon unacquainted, though it was with some surprise that I found in his paper no reference to the phenomenon.

For as it happens, xenia is not at all a rare occurrence with certain varieties of peas; though in them, as I believe is generally the case with this phenomenon, it is highly irregular in its manifestations, being doubtless dependent on slight differences of conditions during ripening.

The coats of peas differ greatly in different varieties, being sometimes thick and white or yellow, sometimes thick and highly pigmented with green or other colours, in both of which cases it may be impossible to judge the cotyledon-colour without peeling off the opaque coat; or the coats may be very thin, colourless and transparent, so that the cotyledon-colour is seen at once. It was such a transparent form that Mendel says he used for his experiments with cotyledon-colour. In order to see xenia a pea with a pigmented seed-coat should be taken as seed-parent, and crossed with a variety having a different cotyledon-colour. There is then a fair chance of seeing this phenomenon, but much still depends on the variety. For example, Fillbasket has green cotyledons and seed-coat green except near the hilar surface. Crossed with Serpette nain blanc (yellow cotyledons and yellow coat) this variety gave three pods with 17 seeds in which the seed-coats were almost full yellow (xenia). Three other pods (25 seeds), similarly produced, showed slight xenia, and one pod with eight seeds showed little or none.

On the other hand Fillbasket fertilised with nain de Bretagne (yellow cotyledons, seed-coats yellow to yellowish green) gave six pods with 39 seeds showing slight xenia, distinct in a few seeds but absent in most.

Examples of xenia produced by the contrary proceeding, namely fertilising a yellow pea with a green, may indubitably occur and I have seen doubtful cases; but as by the nature of the case these are negative phenomena, i.e. the seed-coat remaining greenish and not going through its normal maturation changes, they must always be equivocal, and would require special confirmation before other causes were excluded.

Lastly, the special change (xenia) Mendel saw in “grey” peas, appearance or increase of purple pigment in the thick coats, following crossing, is common but also irregular.

If a transparent coated form be taken as seed-parent there is no appreciable xenia, so far as I know, and such a phenomenon would certainly be paradoxical82.

In this connection it is interesting to observe that Giltay, whom Professor Weldon quotes as having obtained purely Mendelian results, got no xenia though searching for it. If the reader goes carefully through Giltay’s numerous cases, he will find, almost without doubt, that none of them were such as produce it. Reading Giant, as Giltay states, has a transparent skin, and the only xenia likely to occur in the other cases would be of the peculiar and uncertain kind seen in using “grey” peas. Professor Weldon notes that Giltay, who evidently worked with extreme care, peeled his seeds before describing them, a course which Professor Weldon, not recognizing the distinction between the varieties with opaque and transparent coats, himself wisely recommends. The coincidence of the peeled seeds giving simple Mendelian results is one which might have alarmed a critic less intrepid than Professor Weldon.

Bearing in mind, then, that the coats of peas may be transparent or opaque; and in the latter case may be variously pigmented, green, grey, reddish, purplish, etc.; that in any of the latter cases there may or may not be xenia; the reader will perceive that to use the statements of an author, whether scientific or lay, to the effect that on crossing varieties he obtained peas of such and such colours without specifying at all whether the coats were transparent or whether the colours he saw were coat- or cotyledon-colours is a proceeding fraught with peculiar and special risks.

(1) Gärtner’s cases. Professor Weldon gives, as exceptions, a series of Gärtner’s observations. Using several varieties, amongst them Pisum sativum macrospermum, a “grey” pea, with coloured flowers and seed-coats83, he obtained results partly Mendelian and partly, as now alleged, contradictory. The latter consist of seeds “dirty yellow” and “yellowish green,” whereas it is suggested they should have been simply yellow.

Now students of this department of natural history will know that these same observations of Gärtner’s, whether rightly or wrongly, have been doing duty for more than half a century as stock illustrations of xenia. In this capacity they have served two generations of naturalists. The ground nowadays may be unfamiliar, but others have travelled it before and recorded their impressions. Darwin, for example, has the following passage84: