In these the wide range of varieties and the large scale in which they are multiplied of course give a greater chance for new varieties. But then the possibilities of crossing are likewise much larger, and apparent changes due to this cause may easily be taken for original mutations.

The rarity of the mutations is often proved by the lapse of time between the introduction of a species and its first sport. Some instances may be given. They afford a proof of the length of the period during which the species remained unaltered, although some of these alterations may be due to a cross with an allied form. _Erythrina Crista-galli_ was introduced about 1770, and produced its first sport in 1884, after more than a century of cultivation. _Begonia semperflorens_ has been cultivated since 1829, and for half a century before it commenced sporting. The same length of time has elapsed [621] between the first culture and the first variation of _Crambe maritima_. Other cases are on record in which the variability exhibited itself much sooner, perhaps within a few years after the original discovery of the species. But such instances seem, as a rule, to be subject to doubt as to the concurrence of hybridization. So for instance the _Iris lortetii_, introduced in the year 1895 from the Lebanon, which produced a white variety from its very first seeds. If by chance the introduced plants were natural hybrids between the species and the white variety, this apparent and rather improbable mutation would find a very simple explanation. The length of the period preceding the first signs of variability is largely, of course, due to divergent methods of culture. Such species as _Erythrina_, which are perennial and only sown on a small scale, should not be expected to show varieties very soon. Annual species, which are cultivated yearly in thousands or even hundreds of thousands of individuals, have a much better chance. Perhaps the observed differences are largely due to this cause.

Monstrosities have, from time to time, given rise to cultivated races.

The c.o.c.ks...o...b..or _Celosia_ is one of the most notorious instances.

Cauliflowers, turnips and varieties of cabbages are recorded by De Candolle to have arisen in [622] culture, more than a century ago, as isolated monstrous individuals. They come true from seed, but show deviations from time to time which seem to be intimately linked with their abnormal characters. Apetalous flowers may be considered as another form of monstrosity, and in _Salpiglossis sinuata_ such a variety without a corolla made its appearance in the year 1892 in the nursery of Vilmorin. It appeared suddenly, yielded a good crop of seed and was constant from the outset, without any sign of vicinism or impurity.

In several cases the origin of a variety is obscure, while the subsequent historical evidence is such as to make an original sudden appearance quite probable. Although these instances offer but indirect evidence, and will sooner or later lose their importance, it seems desirable to lay some stress on them here, because most of these cases are very obvious and more striking than purely historical facts. Sterile varieties belong to this heading. Sometimes they bear fruit without kernels, sometimes flowers without s.e.xual organs, or even no flowers at all. Instances have been given in the lecture on retrograde varieties; they are ordinarily a.s.sumed to have originated by a leap, because it is not quite clear how a loss of the capacity for the formation of seeds could have been slowly acc.u.mulated [623] in preceding generations. An interesting case is afforded by a sterile variety of corn, which originated some time ago in my own pedigree-cultures made for another purpose, and which had begun with an ear of 1886. The first generation from the original seeds showed nothing particular, but the second at once produced quite a number of sterile plants. The sterility was caused by the total lack of branches, including those bearing the pistillate flowers. The terminal spikes themselves were reduced to naked spindles, without branches, without flowers and even almost without bracts.

In some individuals, however, this negative character was seen to give way at the tip, showing a few small naked branches. Of course it was impossible to propagate this curious form, but my observations showed that it sprang into existence from known ancestors by a single step or sudden leap. This leap, however, was not confined to a single specimen; on the contrary it affected 40 plants out of a culture of 340 individuals. The same phenomenon was repeated from the seeds of the normal plants in the following year, but afterwards the monstrosity disappeared.

The Italian poplar affords another instance. It is considered by some authors as a distinct species, _Populus italica_, and by others as a [624] broom-like variety of the _Populus nigra_, from which it is distinguished by its erect branches and other characters of minor importance. It is often called the pyramidal or fastigiate poplar. Its origin is absolutely unknown and it occurs only in the cultivated state.

In Italy it seems to have been cultivated from the earliest historical times, but it was not introduced into other countries till the eighteenth century. In 1749 it was brought into France, and in 1758 into England, and to day it may be seen along roads throughout central Europe and in a large part of Asia. But the most curious fact is that it is only observed in staminate specimens; pistillate trees have not been found, although often sought for. This circ.u.mstance makes it very probable that the origin of the broom-like poplar was a sudden mutation, producing only one individual. This being staminate, it has been propagated exclusively by cuttings. It is to be admitted, however, that no material evidence is at hand to prove that it is not an original wild species, the pistillate form of which has been lost by vegetative multiplication. One form only of many dioecious plants is to be found in cultivation, as, for instance some South American species of _Ribes_.

Total lack of historical evidence concerning [625] the origin of a variety has sometimes been considered as sufficient proof of a sudden origin. The best known instance is that of the renowned cactus-dahlia with its recurved instead of incurved ray-florets. It was introduced from Mexico into the Netherlands by Van den Berg of Jutphaas, under the following remarkable circ.u.mstances. In the autumn of 1872 one of his friends had sent him a small case, containing seeds, bulbs and roots from Mexico. From one of these roots a _Dahlia_ shoot developed. It was cultivated with great care and bloomed next year. It surprised all who saw it by the unexpected peculiarity of its large rich crimson flowers, the rays of which were reversed tubular. The margins of the narrow rays were curved backwards, showing the bright color of the upper surface. It was a very showy novelty, rapidly multiplied by cuttings, and was soon introduced into commerce. It has since been crossed with nearly all other available varieties of the _Dahlia_, giving a large and rich group of forms, bound together by the curious curling of the petals. It has never been observed to grow in Mexico, either wild or in gardens, and thus the introduced individual has come to be considered as the first of its race.

I have already mentioned that the rapid production of large numbers of new varieties, by [626] means of the crossing of the offspring of a single mutant with previously existing sorts, is a very common feature in horticultural practice. It warns us that only a small part of the novelties introduced yearly are due to real mutations. Further instances of novelties with such a common origin are the purple-leaved dahlias, the gooseberries without p.r.i.c.kles, the double petunias, erect gloxinias and many others. Acc.u.mulation of characters, acquired in different races of a species, may easily be effected in this way; in fact it is one of the important factors in the breeding of horticultural novelties.

I have alluded more than once in this lecture to the question, whether it is probable that mutations occur in one individual or in more. The common belief among horticulturists is that, as a rule, they appear in a single plant. This belief is so widespread that whenever a novelty is seen for the first time in two or more specimens it is at once suggested that it might have originated and been overlooked in a previous generation. Not caring to confess a lack of close observation, the number of mutants in such cases is usually kept secret. At least this statement has been made to me by some of the horticulturists at Erfurt, whom I visited some years ago in order to learn as much as [627]

possible about the methods of production of their novelties. Hence it is simply impossible to decide the question on the basis of the experience of the breeders. Even in the case of the same novelty arising in sundry varieties of the same species, the question as to common origin, by means of crossing, is often hard to decide, as for instance in moss-roses and nectarines. On the other hand, instances are on record where the same novelty has appeared at different times, often at long intervals. Such is the case with the b.u.t.terfly-cyclamen, a form with wide-spreading petals which originated in Martin's nursery in England.

The first time it was seen it was thought to be of no value, and was thrown away, but when appearing for a second time it was multiplied and eventually placed on the market. Other varieties of _Cyclamen_, as for instance the crested forms, are also known to have originated repeatedly.

In concluding this series of examples of horticultural mutations, I might mention two cases, which have occurred in my own experimental garden. The first refers to a tubular _Dahlia_. It has ray-florets, the ligules of which have their margins grown together so as to form tubes, with the outer surface corresponding to the pale under-surface of the corolla.

This novelty originated in a single plant in a [628] culture from the seed of the dwarf variety "Jules Chretien." The seeds were taken from introduced plants in my garden, and as the sport has no ornamental value it is uncertain whether this was the first instance or whether it had previously occurred in the nursery at Lyons, from whence the bulbs were secured. Afterwards it proved true from seed, but was very variable, exhibiting rather the features of an ever-sporting variety.

Another novelty was seen the first time in several individuals. It was a pink sport of the European cranesbill, _Geranium pratense_. It arose quite unexpectedly in the summer of 1902 from a striped variety of the blue species. It was seen in seven specimens out of a lot of about a hundred plants. This strain was introduced into my garden in 1897, when I bought two plants under the name of _Geranium pratense alb.u.m_, which however proved to belong to the striped variety. From their seeds I sowed in 1898 a first generation, of which a hundred plants flowered the next year, and from their seeds I sowed in 1900 the lot which produced the sport. Neither the introduced plants nor their offspring had exhibited the least sign of a color-variation, besides the blue and white stripes. Hence it is very probable that my novelty was a true first mutation, the more probably [629] so since a pink variety would without doubt have a certain horticultural value and would have been preserved if it had occurred. But as far as I have been able to ascertain, it is as yet unknown, nor has it been described until today.

Summing up the results of this long, though very incomplete, list of horticultural novelties with a more or less well-known origin, we see that sudden appearances are the rule. Having once sprung into existence the new varieties are ordinarily constant, except as affected by vicinism. Details concerning the process are mostly unavailable or at least are of very doubtful value. And to this it should be added that really progressive mutations have hardly been observed in horticulture.

Hence the theoretical value of the facts is far less than might have been expected.

[630]

LECTURE XXII

SYSTEMATIC ATAVISM

The steady cooperation of progression and retrogression is one of the important principles of organic evolution. I have dwelt upon this point more than once in previous lectures. I have tried to show that both in the more important lines of the general pedigree of the vegetable kingdom, and in the numerous lateral branches ending in the genera and species within the families, progression and retrogression are nearly always at work together. Your attention has been directed to the monocotyledons as an example, where retrogression is everywhere so active that it can almost be said to be the prevailing movement.

Reduction in the vegetative and generative organs, in the anatomical structure and growth of the stems, and in sundry other ways is the method by which the monocotyledons have originated as a group from their supposed ancestors among the lower dicotyledonous families.

Retrogression is the leading idea in the larger families of the group, [631] as for instance in the aroids and the gra.s.ses. Retrograde evolution is also typical in the highest and most highly differentiated family of the monocotyledons, the orchids, which have but one or two stamens. In the second place I have had occasion more than once to a.s.sert that retrogression, though seemingly consisting in the disappearance of some quality, need not, as a rule, be considered as a complete loss. Quite on the contrary, it is very probable that real losses are extremely rare, if not wholly lacking. Ordinarily the loss is only apparent, the capacity becomes inactive only, but is not destroyed.

The character has become latent, as it is commonly stated, and therefore may return to activity and to the full display of its peculiarity, whenever occasion offers.

Such a return to activity was formerly called atavism. But as we have seen, when dealing with the phenomena of latency at large, sundry cases of latency are to be distinguished, in order to get a clear insight into these difficult processes.

So it is with atavism, too. If any plant reverts to a known ancestor, we have a positive and simple case. But ancestors with alternate specific marks are as a rule neither historically nor experimentally manifest.

They are only reputed to be such, and the presumption rests [632] upon the systematic affinity between the derivative species and its nearest probable allies. Such reversions are now to be examined at some length and may be adequately treated under the head of systematic atavism. To this form of atavism pertain, on the basis of our definition, those phenomena by which species a.s.sume one or more characters of allies, from which they are understood to have descended by the loss of the character under discussion. The phenomena themselves consist in the production of anomalies and varieties, and as the genetic relation of the latter is often hardly beyond doubt, the anomalies seem to afford the best instances for the study of systematic atavism. This study has for its chief aim the demonstration of the presence of the latent characters, and to show that they return to activity suddenly and not by a slow and gradual recovery of the former features. It supports the a.s.sertion that the visible elementary characters are essentially an external display of qualities carried by the bearers of heredity, and that these bearers are separate ent.i.ties, which may be mingled together, but are not fused into a chaotic primitive life-substance. Systematic atavism by this means leads us to a closer examination of the internal and concealed causes, which rule the affinities and divergencies of [633] allied species. It brings before us and emphasizes the importance of the conception of the so-called unit-characters.

The primrose will serve as an example. In the second lecture we have seen that the old species of Linnaeus, the _Primula veris_, was split up by Jacquin into three smaller ones, which are called _P. officinalis_, _P. elatior_ and _P. acaulis_. From this systematic treatment we can infer that these three forms are a.s.sumed to be derived from a common ancestor. Now two of them bear their flowers in bracted whorls, condensed into umbels at the summits of a scape. The scapes themselves are inserted in the axils of the basal leaves, and produce the flowers above them. In the third species, _Primula acaulis_, this scape is lacking and the flowers are inserted singly in the axils on long slender stalks. For this reason the species is called acaulescent, indicating that it has no other stem than the subterranean rootstock. But on closer inspection we observe that the flower stalks are combined into little groups, each group occupying the aril of one of the basal leaves. This fact at once points to an a.n.a.logy with the umbellate allies, and induces us to examine the insertion of the flowers more critically. In doing so we find that they are united at their base so as to const.i.tute a sessile umbel. [634] The scapes are not absolutely lacking, but only reduced to almost invisible rudiments.

Relying upon this conclusion we infer that all of the three elementary species have umbels, some pedunculate and the others not. On this point they agree with the majority of the allied species in the genus and in other genera, as for instance in _Androsace_. Hence the conclusion that the common ancestors were perennial plants with a rootstock bearing their flowers in umbels or whorls on scapes. Lacking in the _Primula veris_, these scapes must obviously have been lost at the time of the evolution of this form.

Proceeding on this line of speculation we at once see that a very adequate opportunity for systematic atavism is offered here. According to our general conception the apparent loss of a scape is no proof of a corresponding internal loss, but might as well be caused simply by the reduction of the scape-growing capacity to a latent or inactive state.

It might be awakened afterwards by some unknown agency, and return to activity.

Now this is exactly what happens from time to time. In Holland the acaulescent primrose is quite a common plant, filling the woods in the spring with thousands of cl.u.s.ters of bright yellow flowers. It is a very uniform type, but in [635] some years it is seen to return to atavistic conditions in some rare individuals. More than once I have observed such cases myself, and found that the variation is only a partial one, producing one or rarely two umbels on the same plant, and liable to fail of repet.i.tion when the varying specimens are transplanted into the garden for further observation. But the fact remains that scapes occur.

The scapes themselves are of varying length, often very short, and seldom long, and their umbels display the involucre of bracts in a manner quite a.n.a.logous to that of the _Primula officinalis_ and _P.

elatior_. To my mind this curious anomaly strongly supports the view of the latent condition of the scape in the acaulescent species, and that such a dormant character must be due to a descent from ancestors with active scapes, seems to be in no need of further reiteration. Returning to activity the scapes at once show a full development, in no way inferior to that of the allied forms, and only unstable in respect to their length.

A second example is afforded by the bracts of the crucifers. This group is easily distinguished by its cruciform petals and the grouping of the flowers into long racemes. In other families each flower of such an inflorescence would be subtended by a bract, according to the [636]

general rule that in the higher plants side branches are situated in the arils of leaves. Bracts are reduced leaves, but the spikes of the cruciferous plants are generally devoid of them. The flower-stalks, with naked bases, seem to arise from the common axis at indefinite points.

Hence the inference that crucifers are an exception to a general rule, and that they must have originated from other types which did comply with this rule, and accordingly were in the possession of floral bracts.

Or, in other words, that the bracts must have been lost during the original evolution of the whole family. This conclusion being accepted, the accidental re-apparition of bracts within the family must be considered as a case of systematic atavism, quite a.n.a.logous to the re-appearance of the scapes in the acaulescent primrose. The systematic importance of this phenomenon, however, is far greater than in the first case, in which we had only to deal with a specific character, while the abolition of the bracts has become a feature of a whole family.

This reversion is observed to take place according to two widely different principles. On one hand, bracts may be met with in a few stray species, a.s.suming the rank of a specific character. On the other hand they may be seen [637] to occur as an anomaly, incompletely developed, often very rare and with all the appearance of an accidental variation, but sometimes so common as to seem nearly normal.

Coming now to particular instances, we may turn our attention in the first place to the genus _Sisymbrium_. This is a group of about 50 species, of wide geographic distribution, among which the hedge mustard (_S. officinalis_) is perhaps the most common of weeds. Two species are reputed to have bracts, _Sisymbrium hirsutum_ and _S. supinum_. Each flower-stalk of their long racemes is situated in the aril of such a bract, and the peculiarity is quite a natural one, corresponding exactly to what is seen in the inflorescence of other families. Besides the _Sisymbrium some six other genera afford similar structures.

_Erucastrum pollichii_ has been already alluded to in a former lecture when dealing with the same problem from another point of view. As previously stated, it is one of the most manifest and most easily accessible examples of a latent character becoming active through systematic atavism. In fact, its bracts are found so often as to be considered by some authors as of quite normal occurrence. Contrasted with those of the above mentioned species of _Sisymbrium_, they are not seen at the base of all the flower [638] stalks, but are limited to the lowermost part of the raceme, adorning a few, often ten or twelve, and rarely more flower-stalks. Moreover they exhibit a feature which is indicative of the presence of an abnormality. They are not all of the same size, but decrease in length from the base of the raceme upward, and finally slowly disappear.

Besides these rare cases there are quite a number of cruciferous species on record, which have been observed to bear bracts. Penzig in his valuable work on teratology gives a list of 33 such genera, many of them repeating the anomaly in more than one species. Ordinary cabbages are perhaps the best known instance, and any unusual abundance of nourishment, or anomalous cause of growth seems to be liable to incite the development of bracts. The hedge garlic or garlic mustard (_Alliaria_), the shepherd's purse, the wormseed or _Erysimum cheiranthoides_ and many others afford instances. In my cultures of Heeger's shepherd's purse, the new species derived at Landau in Germany from the common shepherd's purse, the anomaly was observed to occur more than once, showing that the mutation, which changed the fruits, had not in the least affected this subordinate anomalous peculiarity. In all these cases the bracts behave as with the Erucastrum, [639] being limited to the base of the spike, and decreasing in size from the lower flowers upward. Connected with these atavistic bracts is a feature of minor importance, which however, by its almost universal accompaniment of the bracts, deserves our attention, as it is indicative of another latent character. As a rule, the bracts are grown together with their axillary flower-stalk. This cohesion is not complete, nor is it always developed in the same degree. Sometimes it extends over a large part of the two organs, leaving only their tips free, but on other occasions it is limited to a small part of the base. But it is very interesting that this same cohesion is to be seen in the shepherd's purse, in the wormseed and in the cabbage, as well as in the case of the _Erucastrum_ and most of the other observed cases of atavistic bracts. This fact suggests the idea of a common origin for these anomalies, and would lead to the hypothesis that the original ancestors of the whole family, before losing the bracts, exhibited this peculiar mode of cohesion.

Bracts and a.n.a.logous organs afford similar cases of systematic atavism in quite a number of other families. Aroids sometimes produce long bracts from various places on their spadix, as may be seen in the cultivated greenhouse species, _Anthurium scherzerianum_. [640] Poppies have been recorded to bear bracts a.n.a.logous to the little scales on the flower-stalks of the pansies, on the middle of their flower stalks. A similar case is shown by the yellow foxglove or _Digitalis parviflora_.

The foxgloves as a rule have naked flower-stalks, without the two little opposite leafy organs seen in so many other instances. The yellow species, however, has been seen to produce such scales from time to time. The honeysuckle genus is, as a rule, devoid of the stipules at the base of the petiole, but _Lonicera etrusca_ has been observed to develop such organs, which were seen to be free in some, but in other specimens were adnate to the base of the leaf, and even connate with those of the opposite leaf.

Other instances could be given proving that bracts and stipules, when systematically lacking, are liable to reappear as anomalies. In doing so, they generally a.s.sume the peculiar characters that would be expected of them by comparison with allied genera in which they are of normal occurrence. There can be no doubt that their absence is due to an apparent loss, resulting from the reduction of a formerly active quality to inactivity. Resuming this effective state, the case attains the value and significance accorded to systematic atavism.

A very curious instance of reduced bracts, developing [641] to unusual size, is afforded by a variety of corn, which is called _Zea Mays cryptosperma_, or _Zea Mays tunicata_. In ordinary corn the kernels are surrounded by small and thin, inconspicuous and membranaceous scales.

Invisible on the integrate spikes, when ripe, they are easily detected by pulling the kernels out. In _cryptosperma_ they are so strongly developed as to completely hide the kernels. Obviously they const.i.tute a case of reversion to the characters of some unknown ancestor, since the corn is the only member of the gra.s.s-family with naked kernels. The var.

_tunicata_, for this same reason, has been considered to be the original wild form, from which the other varieties of corn have originated. But as no historical evidence on this point is at hand, we must leave it as it is, notwithstanding the high degree of attractiveness attached to the suggestion.

The horsetail-family may be taken as a further support of our a.s.sertion.

Some species have stems of two kinds, the fertile being brownish and appearing in early spring before the green or sterile ones. In others the stems are all alike, green and crowned with a conelike spike of sporangia-bearing scales. Manifestly the dimorphous cases are to be considered as the younger ones, partly because they are obvious exceptions to the common rule, and [642] partly because the division of labor is indicative of a higher degree of evolution. But sometimes these dimorphic species are seen to revert to the primary condition, developing a fertile cone at the summit of the green summer-stem. I have had the opportunity of collecting an instance of this anomaly on the tall _Equisetum telmateja_ in Switzerland, and other cases are on record in teratological literature. It is an obvious example of systematic atavism, occurring suddenly and with the full development of all the qualities needed for the normal production of sporangia and spores. All of these must be concealed in a latent condition within the young tissues of the green stems.

More than once I have had occasion to deal with the phenomenon of torsions, as exhibited by the teasels and some other plants. This anomaly has been shown to be a.n.a.logous to the cases described as double adaptations. The capacity of evolving antagonistic characters is prominent in both. The antagonists are a.s.sumed to lie quietly together while inactive. But as soon as evolution calls them into activity they become mutually exclusive, because only one of them can come to full display in the same organ. External influences decide which of the two becomes dominant and which remains dormant. This decision must take place separately [643] for each stem and each branch, but as a rule, the stronger ages are more liable to furnish anomalies than the weaker.

Exactly the same thing is true of double adaptations. Every bud of the water-persicaria may develop either into an erect or into a floating stem, according as it is surrounded by water or by relatively dry soil.

In other cases utility is often less manifest, but some use may either be proved, or shown to be very probable. At all events the term adaptation includes the idea of utility, and obviously useless contrivances could hardly be brought under the same head.

We have also dealt with the question of heredity. It is obvious that from the flowers of the floating and erect stems of the water-persicaria seeds will result, each capable of yielding both forms. Quite the same thing was the case with the teasels. Some 40% of the progeny produce beautifully twisted stems, but whether the seed was saved from the most completely twisted specimens or from the straight plants of the race was of no importance.

This phenomenon of twisting may now be considered from quite another point of view. It is a case of systematic atavism, or of the reacquirement of some ancient and long-lost quality. This quality is the alternate position of [644] the leaves, which has been replaced in the teasel family by a grouping in pairs. In order to prove the validity of this a.s.sertion, it will be necessary to discuss two points separately, viz.: relative positions of the leaves, and the manner in which the alternate position causes the stems to become twisted.

Leaves are affixed to their stems and branches in various ways. Among them one is of wide occurrence throughout the whole realm of the higher plants, while all the others are more rare. Moreover these subordinate arrangements are, as a rule, confined to definite systematic groups.

Such groups may be large, as for instance, the monocotyledons, that have their leaves arranged in two opposite rows in many families, or small, as genera or subdivisions of genera. Apart from these special cases the main stem and the greater part of the branches of the pedigree of the higher plants exhibit a spiral condition or a screw arrangement, all leaves being inserted at different points and on different sides of the stem. This condition is a.s.sumed to be the original one, from which the more specialized types have been derived. As is usual with characters in general, it is seen to vary around an average, the spiral becoming narrower and looser. A narrow spiral condenses the leaves, while a [645]

loose one disperses them. According to such fluctuating deviations the number of leaves, inserted upon a given number of spiral circuits, is different in different species. In a vast majority of cases 13 leaves are found on 5 circuits, and as we have only to deal with this proportion in the teasels we will not consider others.

In the teasels this screw-arrangement has disappeared, and has been replaced by a decussate grouping. The leaves are combined into pairs, each pair occupying the opposite sides of one node. The succeeding pairs alternate with one another, so as to place their leaves at right angles.

The leaves are thus arranged on the whole stem in four equidistant rows.