Species and Varieties, Their Origin by Mutation - LightNovelsOnl.com
You're reading novel online at LightNovelsOnl.com. Please use the follow button to get notifications about your favorite novels and its latest chapters so you can come back anytime and won't miss anything.
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 const.i.tute 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, const.i.tuting 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 a.s.sumed 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 a.s.sertion 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 repet.i.tion 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 const.i.tute 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 s.p.a.ce, 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 pa.s.sed 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 const.i.tute 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] a.s.suming 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 _Lycopersic.u.m 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 const.i.tute 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 _Lycopersic.u.m 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 _Lycopersic.u.m_ 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.
Of late the same mutation has occurred in the garden of C.A. White at Was.h.i.+ngton. The parent form in this case was the "Acme," of the ordinary weak and spreading habit of growth. It is known as one of the best and most stable of the varieties and was grown by Mr. White for many years, and had not given any sign of a tendency towards change. Seeds from some of the best plants in 1899 were sown the following spring, and the young seedlings unexpectedly exhibited a marked difference from their parents.
From the very outset they were more strong and erect, more compact and of a darker green than the "Acme." When they reached the fruiting stage they had developed into typical representatives of the _Lycopersic.u.m solanopsis_ or upright division. The whole lot of plants comprised only some 30 specimens, and this number, of course, is too small to base far-reaching conclusions upon. But all of the lot showed this type, no true "Acme" being seen among them. The fruit differed in flavor, consistency and color from that of the parent, and it also ripened earlier than the latter. No seed was saved from [657] these plants, but the following year the "Acme" was sown again and found true to its type.
Seeds saved from this generation in 1900 have, however, repeated the mutation, giving rise to exactly the same new upright form in 1901. This was called by its originator "The Was.h.i.+ngton." Seeds from this second mutation were kindly sent to me by Mr. White, and proved true to their type when sown in my garden.
Obviously it is to be a.s.sumed in the case of the tomatoes as well as in instances from other genera cited, that characters of ancestors, which are not displayed in their progeny, have not been entirely lost, but are still present, though in a latent condition. They may resume their activity unexpectedly, and at once develop all the features which they formerly had borne.
Latency, from this point of view, must be one of the most common things in nature. All organisms are to be considered as internally formed of a host of units, partly active and partly inactive. Extremely minute and almost inconceivably numerous, these units must have their material representatives within the most intimate parts of the cells.
[658]
LECTURE XXIII
TAXONOMIC ANOMALIES
The theory of descent is founded mainly on comparative studies, which have the advantage of affording a broad base and the convincing effect of concurrent evidence brought together from widely different sources.
The theory of mutation on the other hand rests directly upon experimental investigations, and facts concerning the actual descent of one form from another are as yet exceedingly rare. It is always difficult to estimate the validity of conclusions drawn from isolated instances selected from the whole range of contingent phenomena, and this is especially true of the present case. Systematic and physiologic facts seem to indicate the existence of universal laws, and it is not probable that the process of production of new species would be different in the various parts of the animal and vegetable kingdoms.
Moreover the principle of unit-characters, the preeminent significance of which has come to be more fully recognized of late, is in full harmony [659] with the theory of sudden mutations. Together these two conceptions go to strengthen the probability of the sudden origin of all specific characters.
Experimental researches are limited in their extent, and the number of cases of direct observation of the process of mutation will probably never become large enough to cover the whole field of the theory of descent. Therefore it will always be necessary to show that the similarity between observed and other cases is such as to lift above all doubt the a.s.sertion of their resulting from the same causes.
Besides the direct comparison of the mutations described in our former lectures, with the a.n.a.logous cases of the horticultural and natural production of species and varieties at large, another way is open to obtain the required proof. It is the study of the phenomena, designated by Casimir de Candolle by the name of taxonomic anomalies. It is the a.s.sertion that characters, which are specific in one case, may be observed to arise as anomalies or as varieties in other instances. If they can be shown to be identical or nearly so in both, it is obviously allowable to a.s.sume the same origin for the specific character and for the anomaly. In other terms, the specific marks may be considered as having originated according to the laws [660] that govern the production of anomalies, and we may a.s.sume them to lie within reach of our experiments. The experimental treatment of the origin of species may also be looked upon as a method within our grasp.
The validity and the significance of these considerations will at once become clear, if we choose a definite example. The broadest and most convincing one appears to me to be afforded by the cohesion of the petals in gamopetalous flowers. According to the current views the families with the petals of their flowers united are regarded as one or two main branches of the whole pedigree of the vegetable kingdom.
Eichler and others a.s.sume them to const.i.tute one branch, and therefore one large subdivision of the system. Bessey, on the other hand, has shown the probability of a separate origin for those groups which have inferior ovaries. Apart from such divergencies the connation of the petals is universally recognized as one of the most important systematic characters.
How may this character have originated? The heath-family or the Ericaceae and their nearest allies are usually considered to be the lowest of the gamopetalous plants. In them the cohesion of the petals is still subject to reversionary exceptions. Such cases of atavism may [661] be observed either as specific marks, or in the way of anomalies.
_Ledum_, _Monotropa_ and _Pyrola_, or the Labrador tea, the Indian pipe and wintergreen are instances of reversionary gamopetalism with free petals. In heaths (_Erica Tetralix_) and in rhododendrons the same deviation is observed to occur from time to time as an anomaly, and even the common _Rhododendron pontic.u.m_ of our gardens has a variety in which the corolla is more or less split. Sometimes it exhibits five free petals, while at other times only one or two are entirely free, the remaining four being incompletely loosened.
Such cases of atavism make it probable that the coherence of the petals has originally arisen by the same method, but by action in the opposite direction. The direct proof of this conclusion is afforded by a curious observation, made by Vilmorin upon the bright and large-flowered garden-poppy, _Papaver bracteatum_. Like all poppies it has four petals, which are free from one another. In the fields of Messrs. Vilmorin, where it is largely cultivated for its seeds, individuals occur from time to time which are anomalous in this respect. They exhibit a tendency to produce connate petals. Their flowers become monopetalous, and the whole strain is designated by the name of _Papaver_ [662]
_bracteatum monopetalum_. Henry de Vilmorin had the kindness to send me some of these plants, and they have flowered in my garden during several years. The anomaly is highly variable. Some flowers are quite normal, exhibiting no sign of connation; others are wholly gamopetalous, the four petals being united from their base to the very margin of the cup formed. In consequence of the broadness of the petals however, this cup is so wide as to be very shallow.
Intermediate states occur, and not infrequently. Sometimes only two or three petals are united, or the connation does not extend the entire length of the petals. These cases are quite a.n.a.logous to the imperfect splitting of the corolla of the rhododendron. Giving free rein to our imagination, we may for a moment a.s.sume the possibility of a new subdivision of the vegetable kingdom, arising from Vilmorin's poppy and having gamopetalous flowers for its chief character. If the character became fixed, so as to lose its present state of variability, such a group of supposit.i.tious gamopetalous plants might be quite a.n.a.logous to the corresponding real gamopetalous families. Hence there can be no objection to the view, that the heaths have arisen in an a.n.a.logous manner from their polypetalous ancestors. Other species of [663] the same genus have also been recorded to produce gamopetalous flowers, as for instance, _Papaver hybridum_, by Hoffmann. Poppies are not the sole example of accidental gamopetaly. Linnaeus observed the same deviation long ago for _Saponaria officinalis_, and since, it has been seen in _Clematis Vitalba_ by Jaeger, in _Peltaria alliacea_ by Schimper, in _Silene annulata_ by Boreau and in other instances. No doubt it is not at all of rare occurrence, and the origin of the present gamopetalous families is to be considered as nothing extraordinary. It is, as a matter of fact, remarkable that it has not taken place in more numerous instances, and the mallows show that such opportunities have been available at least more than once.
Other instances of taxonomic anomalies are afforded by leaves. Many genera, the species of which mainly bear pinnate or palmate leaves, have stray types with undivided leaves. Among the brambles, _Rubus odoratus_ and _R. flexuosus_ may be cited, among the aralias, _Aralia cra.s.sifolia_ and _A. papyrifera_, and among the jasmines, the deliciously scented sambac (_Jasminum Sambac_). But the most curious instance is that of the telegraph-plant, or _Desmodium gyrans_, each complete leaf of which consists of a large terminal leaflet and two little lateral ones. These latter keep up, [664] night and day, an irregular jerking movement, which has been compared to the movements of a semaph.o.r.e. _Desmodium_ is a papilionaceous plant and closely allied to the genus _Hedysarum_, which has pinnate leaves with numerous pairs of leaflets. Its place in the system leaves no doubt concerning its origin from pinnate-leaved ancestors. At the time of its origination its leaves must have become reduced as to the number of the blades, while the size of the terminal leaflet was correspondingly increased.
It might seem difficult to imagine this great change taking place suddenly. However, we are compelled to familiarize ourselves with such hypothetical a.s.sumptions. Strange as they may seem to those who are accustomed to the conception of continuous slow improvements, they are nevertheless in complete agreement with what really occurs. Fortunately the direct proof of this a.s.sertion can be given, and in a case which is narrowly related, and quite parallel to that of the _Desmodium_, since it affects a plant of the same family. It is the case of the monophyllous variety of the b.a.s.t.a.r.d-acacia or _Robinia Pseud-Acacia_. In a previous lecture we have seen that it originated suddenly in a French nursery in the year 1855. It can be propagated by seed, and exhibits a curious degree [665] of variability of its leaves. In some instances these are one-bladed, the blade reaching a length of 15 cm., and hardly resembling those of the common b.a.s.t.a.r.d-acacia. Other leaves produce one or two small leaflets at the base of the large terminal one, and by this contrivance are seen to be very similar to those of the _Desmodium_, repeating its chief characters nearly exactly, and only differing somewhat in the relative size of the various parts. Lastly real intermediates are seen between the monophyllous and the pinnate types.
As far as I have been able to ascertain, these are produced on weak twigs under unfavorable conditions; the size of the terminal leaflet decreases and the number of the lateral blades increases, showing thereby the presence of the original pinnate type in a latent condition.
The sudden origin of this "one-leaved" acacia in a nursery may be taken as a prototype of the ancient origin of _Desmodium_. Of course the comparison only relates to a single character, and the movements of the leaflets are not affected by it. But the monophylly, or rather the size of the terminal blade and the reduction of the lateral ones, may be held to be sufficiently ill.u.s.trated by the b.a.s.t.a.r.d-acacia. It is worth while to state, that a.n.a.logous varieties have also arisen in other genera. The "one-leaved" [666] strawberry has already been referred to. It originated from the ordinary type in Norway and at Paris. The walnut likewise, has its monophyllous variety. It was mentioned for the first time as a cultivated tree about 1864, but its origin is unknown. A similar variety of the walnut, with "one-bladed" leaves but of varying shapes, was found wild in a forest near Dieppe in France some years ago, and appeared to be due to a sudden mutation.
Something more is known concerning the "one-bladed" ashes, varieties of which are often seen in our parks and gardens. The common form has broad and deeply serrate leaves, which are far more rounded than the leaflets of the ordinary ash. The majority of the leaves are simple, but some produce one or two smaller leaflets at their base, closely corresponding in this respect to the variations of the "one-bladed" b.a.s.t.a.r.d-acacia, and evidently indicating the same latent and atavistic character. In some instances this a.n.a.logy goes still further, and incompletely pinnate leaves are produced with two or more pairs of leaflets. Besides this variable type another has been described by Willdenow. It has single leaves exclusively, never producing smaller lateral leaflets, and it is said to be absolutely constant from seed, while the more variable types [667] seem to be also more inconstant when propagated s.e.xually. The difference is so striking and affords such a reliable feature that Koch proposed to make two distinct varieties of them, calling the pure type _Fraxinus excelsior monophylla_, and the varying trees _F. excelsior exheterophylla_. Some writers, and among them Willdenow, have preferred to separate the "one-leaved" forms from the species, and to call them _Fraxinus simplicifolia_.
According to Smith and to Loudon, the "one-leaved" ashes are found wild in different districts in-England. Intermediate forms have not been recorded from these localities. This mode of origin is that already detailed for the laciniate varieties of alders and so many other trees.
Hence it may be a.s.sumed that the "one-leaved" ashes have sprung suddenly but frequently from the original pinnate species. The pure type of Willdenow should, in this case, be considered as due to a slightly different mutation, perhaps as a pure retrograde variety, while the varying strains may only be eversporting forms. This would likewise explain part of their observed inconstancy.
In this respect the historic dates, as collected by Kors.h.i.+nsky, are not very convincing. Vicinism has of course, almost never been excluded, and part of the multiformity of the offspring [668] must obviously be due to this most universal agency. Indirect vicinism also plays some part, and probably affords the explanation of some reputed mutative productions of the variety. So, for instance, in the case of Sinning, who after sowing the seeds of the common ash, got as large a proportion as 2% of monophyllous trees in a culture of some thousand plants. It is probable that his seeds were taken partly from normal plants, and partly from hybrids between the normal and the "one-bladed" type, a.s.suming that these hybrids have pinnate leaves like their specific parent, and bear the characters of the other parent only in a latent condition.
Our third example relates to peltate leaves. They have the stalk inserted in the middle of the blade, a contrivance produced by the connation of the two basal lobes. The water-lilies are a well known instance, exhibiting sagittate leaves in the juvenile stage and changing in many species, into nearly circular peltate forms, of which _Victoria regia_ is a very good example, although its younger stages do not always excite all the interest they deserve. The Indian cress (_Tropaeolum_), the marsh pennywort or _Hydrocotyle_, and many other instances could be quoted. Sometimes the peltate leaves are not at all orbicular, but are elongated, oblong or elliptic, and with only the lobes [669] at the base united. The lemon-scented _Eucalyptus citriodora_ is one of the most widely known cases. In other instances the peltate leaves become more or less hollow, const.i.tuting broad ascidia as in the case of the cra.s.sulaceous genus _Umbilicus_.
This connation of the basal lobes is universally considered as a good and normal specific character. Nevertheless it has its manifest a.n.a.logy in the realm of the anomalies. This is the pitcher or ascidium. On some trees it is of quite common occurrence, as on the lime-tree (_Tilia parvifolia_) and the magnolia (_Magnolia obovata_ and its hybrids). It is probable that both these forms have varieties with, and others without, ascidia. Of the lime-tree, instances are known of single trees which produce hundreds of such anomalous leaves yearly, and one such a tree is growing in the neighborhood of Amsterdam at Lage Vuursche. I have alluded to these cases more than once, but on this occasion a closer inspection of the structure of the ascidium is required. For this purpose we may take the lime-tree as an example. Take the shape of the normal leaves in the first place. These are cordate at their base and mainly inequilateral, but the general shape varies to a considerable extent. This variation is closely related to the position of the leaves on the twigs, and shows [670] distinct indications of complying with the general law of periodicity. The first leaves are smaller, with more rounded lobes, the subsequent leaves attain a larger size, and their lobes slightly change their forms. In the first leaves the lobes are so broad as to touch one another along a large part of their margins, but in organs formed later this contact gradually diminishes and the typical leaves have the lobes widely separated. Now it is easily understood that the contact or the separation of the lobes must play a part in the construction of the ascidia, as soon as the margins grow together.
Leaves which touch one-another, may be affected by the connation without any further malformation. They remain flat, become peltate and exhibit a shape which in some way holds a middle position between the pennyworts and the lemon-scented eucalyptus. Here we have the repet.i.tion of the specific characters of these plants by the anomaly of another. Whenever the margins are not in contact, and become connate, notwithstanding their separation, the blade must be folded together in some slight degree, in order to produce the required contact. This is the origin of the ascidium. It is quite superfluous to insist upon the fact that their width or narrowness must depend upon the corresponding normal form. The more distant the [671] lobes, the deeper the ascidium will become. It should be added that this explanation of the different shapes of ascidia is of general validity.
Ascidia of the snake-plantain or _Plantago lanceolata_ are narrow tubes, because the leaves are oblong or lanceolate, while those of the broad leaved species of arrowhead, as for instance, the _Sagittaria j.a.ponica_, are of a conical shape.