Chapter 36

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 analogous 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 analogous 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 assume 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 constitute a case of reversion to the characters of some unknown ancestor, since the corn is the only member of the grass-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 assertion. 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 analogous to the cases described as double adaptations. The capacity of evolving antagonistic characters is prominent in both. The antagonists are assumed 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 assertion, 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 assumed 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.

On the normal stem of a teasel the two members of a pair are tied to one another in a comparatively complicated way. The leaves are broadly sessile and their bases are united so as to constitute a sort of cup. The margins of these cups are bent upward, thereby enabling them to hold water, and after a rainfall they may be seen filled to the brim. It is believed that these little reservoirs are useful to the plant during the flowering period, because they keep the ants away from the honey. Considering the internal structure of the stem at the base of these cups we find that the vascular bundles of the two opposite leaves are strongly connected [646] with one another, constituting a ring which narrowly surrounds the stem, and which would impede an increase in thickness, if such were in the nature of the plant. But since the stems end their existence during the summer of their development, this structure is of no real harm.

The grouping of the leaves in alternate pairs may be seen within the bud as well as on the adult stems. In order to do this, it is necessary to make transverse sections through the heart of the rosette of the leaves of the first year. If cut through the base, the pair exhibit connate wings, corresponding to the water-cups; if cut above these, the leaves seem to be free from one another.

In order to compare the position of leaves of the twisted plants with this normal arrangement, the best way is to make a corresponding section through the heart of the rosette of the first year. It is not necessary to make a microscopic preparation. In the fall the changed disposition may at once be seen to affect the central leaves of the group. All the rosettes of the whole race commence with opposite leaves; those that are to produce straight stems remain in this condition, but the preparation for twisting begins at the end of the first year as shown by a special arrangement of the leaves. This [647] disposition may then be seen to extend to the very center of the rosette, by use of microscopical sections. Examining sections made in the spring, the original arrangement of the leaves of the stem is observed to continue until the beginning of the growth of the shoot. It is easy to estimate the number of leaves corresponding to a given number of spiral circuits in these sections and the proportion is found to indicate 13 leaves on 5 turns. These figures are the same as those given above for the ordinary arrangement of alternate leaves in the main lines of the pedigree of the vegetable kingdom.

Leaving aside for the moment the subsequent changes of this spiral arrangement, it becomes at once clear that here we have a case of systematic atavism. The twisted teasels lose their decussation, but in doing so the leaves are not left in a disorderly dispersion, but a distinct new arrangement takes its place, which is to be assumed as the normal one for the ancestors of the teasel family. The case is to be considered as one of atavism. Obviously no other explanation is possible, than the supposition that the 5-13 spiral is still latent, though not displayed by the teasels. But in the very moment when the faculty of decussation disappears, it resumes its place, and becomes [648] as prominent as it must once have been in the ancestors, and is still in that part of their offspring, which has not become changed in this respect. Thus the proof of our assertion of systematic atavism is, in this case, not obtained by the inspection of the adult, but by the investigation of the conditions in an early stage. It remains to be explained how the twisting may finally be caused by this incipient grouping of the leaves. Before doing so, it may be as well to state that the case of the teasel is not an isolated one, and that the same conclusions are supported by the valerian, and a large number of other examples. In early spring some rosettes show a special condition of the leaves, indicating thereby at once their atavism and their tendency to become twisted as soon as they begin to expand. The Sweet William or _Dianthus barbatus_ affords another instance; it is very interesting because a twisted race is available, which may produce thousands of instances developed in all imaginable degrees, in a single lot of plants. _Viscaria oculata_ is another instance belonging to the same family.

The bedstraw (_Galium_) also includes many species which from time to time produce twisted stems. I have found them myself in Holland on _Galium verum_ and _G. Aparine_. Both seem [649] to be of rare occurrence, as I have not succeeded in getting any repetition by prolonged culture.

Species, which generally bear their leaves in whorls, are also subjected to casual atavisms of this kind, as for instance the tall European horsetail, _Equisetum Telmateja_, which occasionally bears cones on its green summer stems. Its whorls are changed on the twisted parts into clearly visible spirals. The ironwood or _Casuarina quadrivalvis_ is sometimes observed to produce the same anomaly on its smaller lateral branches.

Coming now to the discussion of the way in which the twisting is the result of the spiral disposition of the leaves, we may consider this arrangement on stems in the adult state. These at once show the spiral line and it is easy to follow this line from the base up to the apex. In the most marked cases it continues without interruption, not rarely however, ending in a whorl of three leaves and a subsequent straight internode, of which there may even be two or three. The spiral exhibits the basal parts of the leaves, with the axillary lateral branches. The direction of the screw is opposed to that of the twisting, and the spiral ribs are seen to cross the line of insertion of the leaves at nearly right angles. On this line the leaves are nearer [650] to one another than would correspond to the original proportion of 5 turns for 13 leaves. In fact, 10 or even 13 leaves may not rarely be counted on a single turn. Or the twist may become so strong locally as to change the spiral into a longitudinal line. On this line all inserted leaves extend themselves in the same direction, resembling an extended flag.

The spiral on the stem is simply the continuation of the spiral line from within the rosettes of the first year. Accordingly it is seen to become gradually less steep at the base. For this reason it must be one and the same with this line, and in extreme youth it must have produced its leaves at the same mutual distances as this line. Transverse sections of the growing summits of the stems support this conclusion.

From these several facts we may infer that the steepness of the spiral line increases on the stem, as it is gradually changed into a screw. Originally 5 turns were needed for 13 leaves, but this number diminishes and 4 or 3 or even 2 turns may take the same number of foliar organs, until the screw itself is changed into a straight line.

This change consists in an unwinding of the whole spiral, and in order to effect this the stem must become wound up in the opposite direction. The winding of the foliar screw must [651] curve the longitudinal ribs. The straighter and steeper the screw becomes, the more the ribs will become twisted. That this happens in the opposite direction is obvious, without further discussion. The twisting is the inevitable consequence of the reversal of the screw.

Two points remain to be dealt with. One is the direct proof of the reversal of the screw, the other the discussion of its cause. The first may be observed by a simple experiment. Of course it proceeds only slowly, but all that is necessary is to mark the position of one of the younger leaves of a growing stem of a twisting individual and to observe the change in its position in a few hours. It will be seen to have turned some way around the stem, and finally may be seen to make a complete revolution in the direction opposite to the screw, and thereby demonstrating the fact of its uncurling.

The cause of this phenomenon is to be sought in the intimate connection of the basal parts of the leaves, which we have detailed above. The fibrovascular strands constitute a strong rope, which is twisted around the stem along the line on which the leaves are inserted. The strengthening of the internodes may stretch this rope to some extent, but it is too strong to be rent asunder. Hence it opposes the normal growth, and the only manner in which the internodes [652] may adjust themselves to the forces which tend to cause their expansion is by straightening the rope. In doing so they may find the required space, by growing out in an unusual direction, bending their axes and twisting the ribs.

To prove the validity of this explanation, a simple experiment may be given. If the fibrovascular rope is the mechanical impediment which hinders the normal growth, we may try the effect of cutting through this rope. By this means the hindrance may at least locally be removed. Now, of course, the operation must be made in an early stage before, or at the beginning of the period of growth, in every case before the uncurling of the rope begins. Wounds made at this time are apt to give rise to malformations, but notwithstanding this difficulty I have succeeded in giving the necessary proof. Stems operated upon become straight where the rope is cut through, though above and under the wounded part they go on twisting in the usual way.

Sometimes the plants themselves succeed in tearing the rope asunder, and long straight internodes divide the twisted stems in two or more parts in a very striking manner. A line of torn leaf-bases connects the two parts of the screw and gives testimony of what has passed within [653] the tissues. At other times the straightening may have taken place directly internal to a leaf, and it is torn and may be seen to be attached to the stem by two distinct bases.

Summing up this description of the hereditary qualities of our twisted teasels and of their mechanical consequences, we may say that the loss of the normal decussation is the cause of all the observed changes. This special adaptation, which places the leaves in alternating pairs, replaced and concealed the old and universal arrangement on a screw line. In disappearing, it leaves the latter free, and according to the rule of systematic atavism, this now becomes active and takes its place. If the fibrovascular connection of the leaf-bases were lost at the same time the stems would grow and become straight and tall. This change however, does not occur, and the bases of the leaves now constitute a continuous rope instead of separate rings, and thereby impede the stretching of the internodes. These in their turn avoid the difficulty by twisting themselves in a direction opposite to that of the spiral of the leaves.

As a last example of systematic atavism I will refer to the reversionary changes, afforded by the tomatoes. Though the culture of this plant is a recent one, it seems to be at present in a state of mutability, producing new strains, or [654] assuming the features of their presumable ancestors. In his work "The Survival of the Unlike," Bailey has given a detailed description of these various types. Moreover, he has closely studied the causes of the changes, and shown the great tendency of the tomatoes to vicinism. By far the larger part of the observed cases of running out of varieties are caused by accidental crosses through the agency of insects. Even improvements are not rarely due to this cause. Besides these common and often unavoidable changes, others of greater importance occur from time to time. Two of them deserve to be mentioned. They are called the "Upright" and the "Mikado" types, and differ as much or even more from their parents than the latter do from any one of their wild congeners. Their characters come true from seed. The "Mikado" race or the _Lycopersicum grandifolium_ (_L. latifolium_) has larger and fewer leaflets than the slender and somewhat flimsy foliage of the common form. Flat or plane blades with decurrent margins constitute another character. This variety, however, does not concern our present discussion. The upright type has stiff and self-sustaining stems and branches, resembling rather a potato-plant than a tomato. Hence the name _Lycopersicum solanopsis_ or _L. validum_, under which it is usually described. [655] The foliage of the plant is so distinct as to yield botanical characters of sufficient importance to justify this specific designation. The leaflets are reduced in numbers and greatly modified, and the flowers in the inflorescence are reduced to two or three. This curious race came in suddenly, without any premonition, and the locality and date of its mutation are still on record. Until some years ago it had not made its appearance for a second time. Obviously it is to be considered as a reversionary form. The limp stems of the common tomatoes are in all respects indicative of the cultivated condition. They cannot hold themselves erect, but must be tied up to supports. The color of the leaves is a paler green than should be expected from a wild plant. Considering other species of the genus _Solanum_, of which the _Lycopersicum_ is a subdivision, the stems are as a rule erect and self-supporting, with some few exceptions. These, however, are special adaptations as shown by the winding stems of the bitter-sweet.

From this discussion we seem justified in concluding that the original appearance of the upright type was of the nature of systematic atavism. It differs however, from the already detailed cases in that it is not a monstrosity, nor an ever-sporting race, but is as constant a form [656] as the best variety or species. Even on this ground it must be considered as a representative of a separate group of instances of the universal rule of systematic reversions.