One of them produced during the ensuing summer one four-bladed and one five-bladed leaf. The seeds were saved separately and sown the following spring and the expected result could soon be seen. Among some 250 individual plants I counted 22 with one or two deviations, and 10 with from three to nine four- or five-bladed leaves. Proportions nearly similar have been observed repeatedly. Better nourished individuals have produced more deviating leaves on one plant, partly owing to the larger number of stems and branches, and poor or average specimens have mostly been without any aberration or with only one or two abnormal leaves. No further improvement could be attained. Quadrifoliolate leaves were always rare, never [360] attaining a number that would put its stamp on a whole bed. I have endeavored to get some six- and seven-bladed crimson clover leaves, but in vain; selection, culture of many hundreds of individuals, manure, and the best possible treatment has not been adequate to produce them. Of course I am quite convinced that a repet.i.tion of my experiment on a far larger scale would yield the desired types, but then only in such rare instances that they would have no influence whatever on the average, or on the improvement of the race.

The eighth generation in the year 1903 has not been noticeably better than the second and third generations after the first selection.

In comparing this statement with the results gained in the experiment with the red clover, the difference is at once striking. In one case a rich variety was isolated, and, by better treatment and sharp methods of selection, was brought up in a few years to its highest pitch of development. In the other case a very weak race was shown to exist, and no amount of work and perseverance was adequate to improve it to any noticeable degree.

I wish to point out that the decision of what is to be expected from deviating specimens may become manifest within one or two generations.

Even the generation grown from the seeds of [361] the first observed aberrant-individuals, if gathered after sufficient isolation during the period of blossoming, may show which type of inheritance is present, whether it is an unpromising half-race, or a richly endowed sporting variety. I have kept such strains repeatedly after the first isolation, and a special case, that of cotyledoneous aberrations, will be dealt with later. The first generation always gave a final decision, provided that a suitable method of cultivation for the species under observation was found at the beginning. This however, is a condition, which it is not at all easy to comply with, when new sorts are introduced into a garden. Especially so when they had been collected in the wild state.

Often one or two years, sometimes more, are necessary to find the proper method of sowing, manuring, transplanting and, other cultural methods satisfactory to the plants. Many wild species require more care and more manure in gardens than the finest garden flowers. And a large number are known to be dependent on very particular conditions of soil.

One of the most curious features of anomalies, which has been learned from acc.u.mulated instances, is the fact that they obey definite laws as to their occurrence on the different parts of the plant. Obviously such laws are [362] not apparent as long as each plant produces only one or two, or, at most, a few instances of the same deviation. On the contrary, any existing regularity must betray itself, as soon as a larger number of instances is produced. A rule of periodicity becomes most clearly manifest in such cases.

This rule is shown by no other race in a more undoubted and evident manner than by the "five-leaved" clover. Evidently the several degrees of deviation, going from three to seven leaflets, may be regarded as responses to different degrees of variation, and their distribution over the stems and branches, or over the whole plant, may be considered as the manifestation of the ever-changing internal tendency to vary.

Considered from this point of view, my plants always showed a definite periodicity in this distribution, which is the same for the whole plant.

Each of them, and each of the larger branches, begin with atavistic leaves or with slight deviations. These are succeeded by greater deviations, but only the strongest axes show as many as seven leaflets on a stalk. This ordinarily does not occur before the height of development is reached, and often only towards its close. Then the deviation diminishes rapidly, returning often to atavistic leaves at the summit of the stem or branch. I give the numbers of the [363] leaves of a branch, in their order from the base to the top. They were as follows:

3. 4. 5. 6. 7. 5. 5. 4.

But this is a selected case, and such regular examples of the expected periodicity are rarely found. Often one or more of the various steps are lacking, or even leaves with smaller numbers may be interspersed among those with larger numbers of leaflets. But while the regularity of the periodicity is in some degree diminished by such occurrences, yet the rule always holds good, when taken broadly. It may be expressed by stating that the bases and apices have on the average fewer leaflets on each leaf than the middle parts of the stem and branches, and that the number of leaflets gradually increases from the base toward a maximum, which is reached in organs on the middle or upper part of the axis, and then diminishes from this toward the apex.

This periodicity is not limited to the stems and branches, considered singly, but also holds good in a comparison made between the branches of a single stem, in regard to their relative places on that stem. So it is also for the whole plant. The first stems, produced by the subterranean axis, ordinarily show only a low maximum deviation: the next succeeding being [364] more divergent and the last ones returning to less differentiated forms.

It is evident that on a given stem the group of deviating leaves will be extended upward and downward, with the increase of the number of these organs. This shows that a stem, or even a plant, promises a higher degree of differentiation if it commences with its aberration earlier.

Hence it becomes possible to discern the most promising individuals in early youth, and this conclusion leads to a very easy and reliable method of selection, which may be expressed simply as follows: the seedlings which commence earliest with the production of four- and five-foliolate leaves are the best and should be selected for the continuance of the race. And it is easily seen that this rule agrees with that given above, and which was followed in my pedigree-culture.

Furthermore it is seen that there is a complete agreement between the law of periodicity and the responses of the deviations to nourishment and other conditions of life. Weak plants only produce low degrees of deviation, the stronger the individual becomes, the higher it reaches in the scale of differentiation, and the more often it develops leaves with five or more blades. Whether weakness or strength are derived from outer causes, or from the internal [365] succession of the periods of life, is evidently of no consequence, and in this way the law of periodicity may be regarded as a special instance of the more general law of response to external conditions.

The validity of this law of periodicity is of course not limited to our "five-leaved" clover. Quite on the contrary it is universal in eversporting varieties. Moreover it may be ascertained and studied in connection with the most widely different morphologic abnormalities, and therefore affords easily accessible material for statistical inquiry. I will now give some further instances, but wish to insist first upon the necessity of an inquiry on a far larger scale, as the evidence as yet is very scanty.

The great celandine (_Chelidonium majus_) has a very curious double variety. Its flowers are simpler and much more variable than in ordinary garden-varieties. The process of doubling consists mainly in a change of stamens into petals. This change is dependent on the season. On each stem the earliest flowers are single. These are succeeded by blossoms with one or two converted stamens, and towards the summer this number increases gradually, attaining 10-11 and in some instances even more altered filaments. Each year the same succession may be seen repeating itself on the stems of [366] the old roots. Double tuberous begonias are ordinarily absolutely sterile throughout the summer, but towards autumn the new flowers become less and less altered, producing some normal stamens and pistils among the majority of metamorphosed organs. From these flowers the seeds are saved. Sometimes similar flowers occur at the beginning of the flowering-period. Double garden-camomiles (_Chrysanthemum inodorum plenissimum_) and many other double varieties of garden-plants among the great family of the composites are very sensitive to external agencies, and their flower-heads are fuller the more favorable the external conditions. Towards the autumn many of them produce fewer and fewer converted heads and often only these are fertile and yield seeds.

Ascidia afford another instance of this periodicity, though ordinarily they are by far too rare to show any regularity in their distribution.

However, it is easy to observe that on lime-trees they prefer the lower parts of each twig, while on magnolias the terminal leaves of the branches are often pitcher-bearing. Ascidia of the white clover have been found in numbers, in my own experiment-garden, but always in the springtime. The thickleaved saxifrage (_Saxifraga cra.s.sifolia_) is often very productive of ascidia, especially in [367] the latter part of the season, and as these organs may be developed to very different degrees, they afford fine material for the study of the law of periodicity. On a garden-cytisus (_Cytisus candicans attleya.n.u.s_) I once had the good fortune to observe a branch with ascidia, which ordinarily are very rare in this species. It had produced seven ascidia in all, each formed by the conversion of one leaflet on the trifoliolate leaves. The first six leaves were dest.i.tute of this malformation and were quite normal. Then followed a group of five leaves, const.i.tuting the maximum of the period.

The first bore one small pitcher-like blade, the second and third, each one highly modified organ, the fourth, two ascidia, and the last, one leaflet with slightly connate margins. The whole upper part of the branch was normal, with the exception of the seventeenth leaf, which showed a slight change in the same direction. All in all, the tendency to produce ascidia increased from the beginning to the tenth leaf, and decreased from this upward.

The European Venus' looking-gla.s.s was observed in my garden to produce some quaternate and some quinate flowers on the same specimens. The quinate were placed at the end of the branches, those with four petals and sepals lower down. The peloric fox-glove shows the [368] highest degree of metamorphy in the terminal flowers of the stem itself, the weaker branches having but little tendency towards the formation of the anomaly. The European pine or _Pinus sylvestris_ ordinarily has two needles in each sheath, but trifoliolate sheaths occur on the stems and stronger branches, where they prefer, as a rule, the upper parts of the single annual shoots. _Camellia j.a.ponica_ is often striped in the fall and during the winter, but when flowering in the spring it returns to the monochromatic type.

Peloric flowers are terminal in some cases, but occur in the lower parts of the flower-spikes in others. Some varieties of gladiolus commence on each spike with more or less double flowers, which, higher up, are replaced by single ones. A wide range of bulbs and perennial garden-plants develop their varietal characters only partly when grown from seed and flowering for the first time. The annual garden-forget-me-not of the Azores (_Myosotis azorica_) has a variety with curiously enlarged flowers, often producing 20 or more corolla-segments in one flower. But this number gradually diminishes as the season advances. It would be quite superfluous to give further proof of the general validity of the law of periodicity in ever-sporting varieties.

[369]

LECTURE XIII

PISTILLODY IN POPPIES

One of the most curious anomalies that may be met with in ornamental garden-plants is the conversion of stamens into pistils. It is neither common nor rare, but in most cases the change is so slight comparatively that it is ordinarily overlooked. In the opium-poppy, on the contrary, it is very showy, and heightens the ornamental effect of the young fruits after the fading of the flowers. Here the central capsule is surrounded by a large crown of metamorphosed stamens.

This peculiarity has attracted the attention both of horticulturists and of botanists. As a rule not all the stamens are changed in this way but only those of the innermost rows. The outer stamens remain normal and fertile, and the flowers, when pollinated with their own pollen, bear as rich a harvest of seeds as other opium-poppies. The change affects both the filament and the anther, the former of which is dilated into a sheath. Within this sheath perfect [370] and more or less numerous ovules may be produced. The anthers become rudimentary and in their place broad leafy flaps are developed, which protrude laterally from the tip and const.i.tute the stigmas. Ordinarily these altered organs are sterile, but in some instances a very small quant.i.ty of seed is produced, and when testing their viability I succeeded in raising a few plants from them.

The same anomaly occurs in other plants. The common wall-flower (_Cheiranthus Cheiri_) and the houseleek (_Sempervivum tectorum_) are the best known instances. Both have repeatedly been described by various investigators. In compiling the literature of this subject it is very interesting to observe the two contrasting views respecting the nature of this anomaly. Some writers, and among them Masters in his "Vegetable Teratology" consider the deviations to be merely accidental. According to them some species are more subject to this anomaly than others, and the houseleek is said to be very p.r.o.ne to this change. Goeppert, Hofmeister and others occasionally found the pistilloid poppies in fields or gardens, and sowed their seeds in order to ascertain whether the accidental peculiarity was inheritable or not. On the other hand De Candolle in his "Prodromus" mentions the pistilloid wall-flowers as a distinct [371] variety, under the name of _Cheiranthus Cheiri gynantherus_, and the a.n.a.logous form of the opium-poppy is not at all an accidental anomaly, but an old true horticultural variety, which can be bought everywhere under the names of _Papaver somniferum monstruosum_ or _polycephalum_. Since it is an annual plant, only the seeds are for sale, and this at once gives a sufficient proof of its heredity. In all cases, where it was met with accidentally by botanists, it is to be a.s.sumed that stray seeds had been casually mixed with those of other varieties, or that the habit had been transmitted by a spontaneous cross.

Wherever opportunity led to experiments on heredity, distinct races were found to be in possession of this quality, while others were not. It is of no use to cultivate large numbers of wall-flowers in the hope of one day seeing the anomaly arise; the only means is to secure the strain from those who have got it. With poppies the various varieties are so often intercrossed by bees, that the appearance of an accidental change may sometimes be produced, and in the houseleek the pistilloid warily seems to be the ordinary one, the normal strain being very rare or perhaps wholly wanting.

Our three ill.u.s.trative examples are good and permanent races, producing their peculiar qualities [372] regularly and abundantly. In this respect they are however very variable and dependent on external circ.u.mstances.

Such a regularity is not met with in other instances. Often pedigree-experiments lead to poor races, betraying their tendency to deviate only from time to time and in rare cases. Such instances const.i.tute what we have called in a former lecture, "half races," and their occurrence indicates that the casual observation of an anomaly is not in itself adequate to give an opinion as to the chance of repet.i.tion in sowing experiments. A large number of species seem to belong to this case, and their names may be found in the above mentioned work by Masters and elsewhere. But no effort has yet been made to separate thoroughly the pistilloid half-races from the corresponding ever-sporting varieties. Some plants are recorded as being more liable to this peculiarity than others.

Stamens are sometimes replaced by open carpels with naked ovules arising from their edges and even from their whole inner surfaces. This may be seen in distinct strains of the cultivated bulbous Begonia, and more rarely in primroses. Here the apex of the carpellary leaf is sometimes drawn out into a long style, terminated by a flattened spatulate stigma.

The pistillody of the stamens is frequently [373] combined with another deviation in the poppies. This is the growing together of some of the altered stamens so as to const.i.tute smaller or larger connate groups.

Often two are united, sometimes three, four or more. Flowers with numerous altered stamens are seldom wholly free from this most undesirable secondary anomaly. I call it undesirable with respect to experiments on the variability of the character. For it may easily be seen that while it is feasible to count the stamens even when converted into pistils, it is not possible when groups of them are more or less intimately united into single bodies. This combination makes all enumeration difficult and inaccurate and often wholly unreliable. In such cases the observation is limited to a computation of the degree of the change, rather than to a strict numerical inquiry. Happily the responses to the experimental influences are so marked and distinct that even this method of describing them has proved to be wholly sufficient.

In extreme instances I have seen all the changed stamens of a flower of the opium-poppy united into a single body, so as to form a close sheath all around the central ovary. Lesser sheaths, surrounding one-half or one-third of the capsule are of course less rarely met with. Leaving this description of the outer appearance [374] of our anomaly, we may now consider it from the double point of view of inheritance and variability.

The fact of inheritance is shown by the experience of many authors, and by the circ.u.mstance already quoted, that the variety has been propagated from seed for more than half a century, and may be obtained from various seed merchants. In respect to the variability, the variety belongs to the ever-sporting group, const.i.tuting a type which is more closely related to the "five-leaved" clover than to the striped flowers or even the double stocks.

It fluctuates around an average type with half filled crowns, going as far as possible in both directions, but never transgressing either limit. It is even doubtful whether the presumable limits are, under ordinary circ.u.mstances, ever reached. Obviously one extreme would be the conversion of all the stamens, and the other the absolute deficiency of any marked tendency to such a change. Both may occur, and will probably be met with from time to time. But they must be extremely rare, since in my own extensive experiments, which were strictly controlled, I never was able to find a single instance of either of them. Some of the outer stamens have always remained unchanged, yielding enough pollen for the artificial pollination of [375] the central ovary, and on the other hand some rudiments of hardened filaments were always left, even if they were reduced to small protuberances on the thalamus of the flower. Between these extremes all grades occur. From single, partially or wholly changed stamens upwards to 150 and over, all steps may be seen. It is a true fluctuating variability. There is an average of between 50 and 100, const.i.tuting a nearly filled crown around the central capsule. Around this average the smaller deviations are most numerous and the larger ones more rare. The inspection of any bed of the variety suffices to show that, taken broadly, the ordinary laws of fluctuating variability are applicable. No counting of the single individuals is required to dispel all doubts on this point.

Moreover all intermediate steps respecting the conversion of the single stamens may nearly always be seen. Rarely all are changed into normal secondary ovaries with a stigma and with a cavity filled with ovules.

Often the stigma is incomplete or even almost wanting, in other instances the ovules are lacking or the cavity itself is only partially developed. Not rarely some stamens are reduced and converted into thin hard stalks, without any appearance of an ovary at their tip. But then the demarcation [376] between them and the thalamus fails, so that they cannot be thrown off when the flower fades away, but remain as small stumps around the base of the more fully converted filaments. This fact would frequently render the enumeration of the altered organs quite unreliable.

For these reasons I have chosen a group of arbitrary stages in order to express the degree of deviation for a given lot of plants. The limits were chosen so as to be sufficiently trustworthy and easy to ascertain.

In each group the members could be counted, and a series of figures was reached by this means which allowed of a further comparison of the competing sets of plants.

It should be stated that in such experiments and especially in the case of such a showy criterion as the pistilloid heads afford after the time of flowering is over, the inspection of the controlling beds at once indicates the result of the experiment. Even a hasty survey is in most cases sufficient to get a definite conclusion. Where this is not the case, the counting of the individuals of the various groups often does not add to the evidence, and the result remains uncertain. On the other hand, the impression made by the groups of plants on the experimenter and on his casual visitors, cannot well be conveyed to the readers of his account by [377] other means than by figures. For this reason the result of the experiments is expressed in this way.

I made six groups. The first includes the cases where the whole circle is reduced to small rudiments. The second shows 1-10 secondary capsules.

The two following const.i.tute half a crown around the central fruit, the third going up to this limit, the fourth going from this limit to a nearly filled circle. Wholly filled circles of secondary capsules without gaps give the two last degrees, the fifth requiring only continuity of the circle, the sixth displaying a large and bright crown all around the central head. The fifth group ordinarily includes from 90-100 altered stamens, while the sixth has from 100-150 of these deviating parts.

In ordinary cultures the third and fourth group, with their interrupted crowns, predominate. Large crowns are rare and flowers which at first sight seem to be wholly normal, occur only under circ.u.mstances definitely known to be unfavorable to growth, and to the development of the anomaly.

Having reached by this means a very simple and easy method of stating the facts shown by equal lots under contrasting influences, we will now make use of it to inquire into the relation [378] of this exceptionally high degree of variability to the inner and outer conditions of life.

As a rule, all experiments show the existence of such a relation.

Unfavorable conditions reduce the numbers of altered stamens, favorable circ.u.mstances raise it to its highest point. This holds true for lots including hundreds of specimens, but also for the sundry heads of one bed, and often for one single plant.

We may compare the terminal flower with those of the lateral branches on a plant, and when no special influences disturb the experiment, the terminal head ordinarily bears the richest crown. If the first has more than 100 metamorphosed parts, the latter have often less than 50 on the same plant. In poor soil, terminal heads are often reduced to 10-20 monstrous organs, and in such cases I found the lateral flowers of the same plants ordinarily with less than 10 altered stamens. In some cases I allowed the branches of the third and fourth degree, in other words, the side twigs of the first branches of my selected plants to grow out and produce flowers in the fall. They were ordinarily weak, sometimes very small, having only 5-9 stigmas on their central fruit. Secondary capsules were not seen on such flowers, even when the experiment was repeated on a [379] somewhat larger scale and during a series of years.

Among the same lot of plants individual differences almost always occur.

They are partly due to inequalities already existing in the seeds, and partly to the diversity of the various parts of the same bed. Some of the plants become stout and have large terminal heads. Others remain very weak, with a slender stem, small leaves and undersized flowers. The height and thickness of the stem, the growth of the foliage and of the axillary buds are the most obvious measures of the individual strength of the plant. The development of the terminal flower and the size of its ovary manifestly depends largely on this individual strength, as may be seen at once by the inspection of any bed of opium-poppies. Now this size of the head can easily be measured, either by its height or circ.u.mference, or by its weight. Moreover we can arrange them into a series according to their size. If we do this with the polycephalous variety, the relation between individual strength and degree of metamorphosis at once becomes manifest. The largest heads have the brightest crowns, and the number of supernumerary carpels diminishes in nearly exact proportion to the size of the fruits. Fruits with less than 50 altered stamens weighed on an average 5 grams, [380] those with 50-100 such organs 7 grams and those with a bright crown 10 grams, the appendices being removed before the weighing. Corresponding results have been reached by the comparison of the height of the capsules with their abnormal surroundings. The degree of development of the monstrosity is shown by this observation to be directly dependent on, and in a sense proportionate to the individual strength of the plant.

The differences between the specimens grown from a single lot of seeds, for instance from the seeds of one self-fertilized capsule are, as I have said, partly due to the divergences which are always present in a bed, even if the utmost care has been taken to make it as uniform as possible. These local differences are ordinarily underrated and overlooked, and it is often considered to be sufficient to cultivate small lots of plants under apparently similar conditions on neighboring beds, to be justified in imputing all the observed deviations of the plants to hereditary inequalities. This of course is true for large lots, whenever the averages only are compared. In smaller experiments the external conditions of the single individuals should always be considered carefully. Lots of one or two square meters suffice for such comparisons, but smaller lots are always subject to chances and [381]

possibilities, which should never be left out of consideration.

Therefore I will now point out some circ.u.mstances, which are ordinarily different on various parts of one and the same bed.

In the first place comes the inequality of the seeds themselves. Some of them will germinate earlier and others later. Those that display their cotyledons on a sunny day will be able to begin at once with the production of organic food. Others appear in bad weather, and will thus be r.e.t.a.r.ded in their development. These effects are of a c.u.mulative nature as the young plants must profit by every hour of sunshine, according to the size of the cotyledons. Any inequality between two young seedlings is apt to be increased by this c.u.mulative effect.

The same holds good for the soil of the bed. It is simply impossible to mix the manure so equally that all individuals receive the same amount of it from the very beginning. I am in the habit of using manures in a dry and pulverized condition, of giving definite quant.i.ties to each square meter, and of taking the utmost care to get equal distribution and mixture with the soil, always being present myself during this most important operation. Nevertheless it is impossible to make the nourishment exactly equal for all the plants of even a small bed.