Species and Varieties: Their Origin by Mutation (1906)
Lectures delivered at the University of California
Professor of Botany in the University of Amsterdam
THE PRODUCTION OF DOUBLE FLOWERS
Mutations occur as often among cultivated plants as among those in the wild state. Garden flowers are known to vary markedly. Much of their variability, however, is due to hybridism, and the combination of characters previously separate has a value for the breeder nearly equal the production of really new qualities. Nevertheless there is no doubt that some new characters appear from time to time.
In a previous lecture we have seen that varietal characters have many features in common. One of them is their frequent recurrence both in the same and in other, often very distantly related, species. This recurrence is an important factor in the choice of the material for an experimental investigation of the nature of mutations.
Some varieties are reputed to occur more often and more readily than others. White-colored varieties, though so very common, seem for the most part to be of ancient date, but only few  have a known origin, however. Without any doubt many of them have been found in a wild state and were introduced into culture. On the other hand double flowers are exceedingly rare in the wild state, and even a slight indication of a tendency towards doubling, the stray petaloid stamens, are only rarely observed growing wild. In cultivation, however, double flowers are of frequent occurrence; hence the conclusion that they have been produced in gardens and nurseries more frequently than perhaps any other type of variety.
In the beginning of my experimental work I cherished the hope of being able to produce a white variety. My experiments, however, have not been successful, and so I have given them up temporarily. Much better chances for a new double variety seemed to exist, and my endeavors in this direction have finally been crowned with success.
For this reason I propose to deal now with the production of double flowers, to inquire what is on record about them in horticultural literature, and to give a full description of the origin thereof in an instance which it was my good fortune to observe in my garden.
Of course the historical part is only a hasty survey of the question and will only give such evidence as may enable us to get an idea of the  chances of success for the experimental worker. In the second half of the seventeenth century (1671), my countryman, Abraham Munting, published a large book on garden plants with many beautiful figures. It is called "Waare Oeffeninge der Planters," or "True Exercises With Plants." The descriptions pertain to ordinary typical species in greater part, but garden varieties receive special attention. Among these a long list of double flowers are to be seen. Double varieties of poppies, liverleaf (Hepatica), wallflowers (Cheiranthus), violets, Caltha, Althaea, Colchicum, and periwinkles (Vinca), and a great many other common flowers were already in cultivation at that time.
Other double forms have been since added. Many have been introduced from Japan, especially the Japanese marigold, Chrysanthemum indicum. Others have been derived from Mexico, as for instance the double zinnias. The single dahlias only seem to have been originally known to the inhabitants of Mexico. They were introduced into Spain at about 1789, and the first double ones were produced in Louvain, Belgium, in 1814. The method of their origin has not been described, and probably escaped the originators themselves. But in historical records we find the curious statement that it took place after three years' work. This indicates  a distinct plan, and the possibility of carrying it to a practical conclusion within a few years' time.
Something more is known about other cases. Garden anemones, Anemone coronaria, are said to have become double in the first half of the last century in an English nursery. The owner, Williamson, observing in his beds a flower with a single broadened stamen, saved its seeds separately, and in the next generations procured beautifully filled flowers. These he afterwards had crossed by bees with a number of colored varieties, and in this way succeeded in producing many new double types of anemone.
The first double petunia is known to have suddenly and accidentally arisen from ordinary seed in a private garden at Lyons about 1855. From this one plant all double races and-varieties have been derived by natural and partly by artificial crosses. Carriere, who reported this fact, added that likewise other species were known at that time to produce new double varieties rapidly. The double fuchsias originated about the same time (1854) and ten years later the range of double varieties of this plant had become so large that Carriere found it impossible to enumerate all of them.
Double carnations seem to be relatively old, double corn-flowers and double blue-bells being  of a later period. A long list could easily be made, to show that during the whole history of horticulture double varieties have arisen from time to time. As far as we can judge, such appearances have been isolated and sudden. Sometimes they sprang into existence in the full display of their beauty, but most commonly they showed themselves for the first time, exhibiting only spare supernumerary petals. Whenever such sports were worked up, a few years sufficed to reach the entire development of the new varietal attribute.
From this superficial survey of historical facts, the inference is forced upon us that the chance of producing a new double variety is good enough to justify the attempt. It has frequently succeeded for practical purposes, why should it not succeed as well for purely scientific investigation? At all events the type recommends itself to the student of nature, both on account of its frequency, and of the apparent insignificance of the first step, combined with the possibility of rapidly working up from this small beginning of one superfluous petal towards the highest degree of duplication.
Compared with the tedious experimental production of the peloric toad-flax, the attempt to produce a double flower has a distinct attraction. The peloric toad-flax is nothing new; the  experiment was only a repetition of what presumably takes place often within the same species. To attempt to produce a double variety we may choose any species, and of course should select one which as yet has not been known to produce double flowers. By doing so we will, if we succeed, produce something new. Of course, it does not matter whether the new variety has an horticultural interest or not, and it seems preferable to choose a wild or little cultivated species, to be quite sure that the variety in question is not already in existence. Finally the prospect of success seems to be enhanced if a species is chosen, the nearest allies of which are known to have produced double flowers.
For these reasons and others I chose for my experiment the corn-marigold, or Chrysanthemum segetum. It is also called the golden cornflower. In the wheat and rye fields of central Europe it associates with the blue-bottle or blue corn-flower. It is sometimes cultivated and the seeds are offered for sale by many nurserymen. It has a cultivated variety, called grandiflorum, which is esteemed for its brilliancy and long succession of golden bloom. This variety has larger flower-heads, surrounded with a fuller border of ray-florets. The species belongs to a genus many species of which have produced  double varieties. One of them is the Japanese marigold, others are the carinatum and the imbricatum species. Nearly allied are quite a number of garden-plants with double flower-heads, among which are the double camomiles.
My attention was first drawn to the structure of the heads and especially to the number of the ray-florets of the corn-marigold. The species appertains to that group of composites which have a head of small tubular florets surrounded by a broad border of rays. These rays, when counted, are observed to occur in definite numbers, which are connected with each other by a formula, known as "the series" of Braun and Schimper. In this formula, which commences with 1 and 2, each number is equal to the sum of the two foregoing figures. Thus 5, 8 and 13 are very frequent occurrences, and the following number, 21, is a most general one for apparently full rays, such as in daisies, camomiles, Arnica and many other wild and cultivated species.
These numbers are not at all constant. They are only the averages, around which the real numbers fluctuate. There may even be an overlapping of the extremes, since the fluctuation around 13 may even go beyond 8 and 21, and so on. But such extremes are only found in stray flowers, occurring on the same  individuals with the lesser degrees of deviation.
Now the marigold averages 13, and the grandiflorum 21 rays. The wild species is pure in this respect, but the garden-variety is not. The seeds which are offered for sale usually contain a mixture of both forms and their hybrids. So I had to isolate the pure types from this mixture and to ascertain their constancy and mutual independency. To this end I isolated from the mixture first the 13-rayed, and afterwards the 21-rayed types. As the marigolds are not sufficiently self-fertile, and are not easily pollinated artificially, it seemed impossible to carry on these two experiments at the same time and in the same garden. I devoted the first three years to the lower form, isolated some individuals with 12-13 rays out of the mixture of 1892 and counted the ray-florets on the terminal head of every plant of the ensuing generation next year. I cultivated and counted in this way above 150 individuals and found an average of exactly 13 with comparatively few individuals displaying 14 or only 12 rays, and with the remainder of the plants grouped symmetrically around this average. I continued the experiment for still another year and found the same group of figures. I was then satisfied as to the purity of the isolated strain. Next year I sowed a new mixture in  order to isolate the reputed pure grandiflorum type. During the beginning of the flowering period I ruthlessly threw away all plants displaying less than 21 rays in the first or terminal head. But this selection was not to be considered as complete, because the 13-rayed race may eventually transgress its boundary and come over to the 21 and more. This made a second selection necessary. On the selected plants all the secondary heads were inspected and their ray-florets counted. Some individuals showed an average of about 13 and were destroyed. Others gave doubtful figures and were likewise eliminated, and only 6 out of a lot of nearly 300 flowering plants reached an average of 21 for all of the flowers.
Our summer is a short one, compared with the long and beautiful summer of California, and it was too late to cut off the faded and the open flowers, and await new ones, which might be purely fertilized after the destruction of all minor plants. So I had to gather the seed from flowers, which might have been partially fertilized by the wrong pollen. This however, is not so great a drawback in selection experiments as might be supposed at first sight. The selection of the following year is sure to eliminate the offspring of such impure parentage.
 A far more important principle is that of the hereditary percentage, already discussed in our lecture on the selection of monstrosities. In our present case it had to be applied only to the six selected plants of 1895. To this end the seeds of each of them were sown separately, the ray-florets of the terminal heads of each of the new generation were counted, and curves and averages were made up for the six groups. Five of them gave proof of still being mixtures and were wholly rejected. The children of the sixth parent, however, formed a group of uniform constitution, all fluctuating around the desired average of 21. All in all the terminal heads of over 1,500 plants have been subjected to the somewhat tedious work of counting their ray-florets. And this not in the laboratory, but in the garden, without cutting them off. Otherwise it would obviously have been impossible to recognize the best plants for preservation. I chose only two plants which in addition recommended themselves by the average number of rays on their secondary heads, sowed their seeds next year separately and compared the numerical constitution of their offspring. Both groups averaged 21 and were distributed very symmetrically around this mean. This result  showed that no further selection could be of any avail, and that I had succeeded in purifying the 21-rayed grandiflorum variety.
It is from this grandiflorum that I have finally produced my double variety. In the year 1896 I selected from among the above quoted 1,500 plants, 500 with terminal heads bearing 21 or more rays. On these I counted the rays of all the secondary heads about the middle of August (1896) and found that they had, as a rule, retrograded to lower figures. On many thousands of heads only two were found having 22 rays. All others had the average number of 21 or even less. I isolated the individual which bore these two heads, allowed them to be fertilized by insects with the pollen of some of the best plants of the same group, but destroyed the remainder.
This single exceptional plant has been the starting point of my double variety. It was not remarkable for its terminal head, which exhibited the average number of rays of the 21-rayed race. Nor was it distinguished by the average figure for all its heads. It was only selected because it was the one plant which had some secondary heads with one ray more than all the others. This indication was very slight, and could not have been detected save by the counting of the rays of thousands of heads.
 But the rarity of the anomaly was exactly the indication wanted, and the same deviation would have had no signification whatever, had it occurred in a group fluctuating symmetrically around the average figure. On the other hand, the observed anomaly was only an indication, and no guarantee of future developments.
Here it should be remarked that the indication alluded to was not the appearance of the expected character of doubling in ever so slight a measure. It was only a guide to be followed in further work. The real character of double flower-heads among composites lies in the production of rays on the disk. No increase of the number of the outer rays can have the same significance. A hasty inspection of double flower heads may convey the idea that all rays are arranged around a little central cluster of disk-florets, the remainder of the original disk-florets but a closer investigation will always reveal the fallacy of this conclusion. Hidden between the inner rays, and covered by them, lie the little tubular and fertile florets everywhere on the disk. They may not be easily seen, but if the supernumerary rays are pulled out, the disk may be seen to bear numerous small florets at intervals. But these intervals are not at all numerous, showing thereby that only a relatively small number of tubes has been  converted into rays. This conversion is obviously the true mark of the doubling, and before traces of it are found, no assertion whatever can be given as to the issue of the pedigree experiment.
Three more years were required before this first, but decisive trace was discovered. During these years I subjected my strain to the same sharp selection as has already been described. The chosen ancestor of the race had flowered in 1896, and the next year I sowed its seeds only. From this generation I chose the one plant with the largest number of rays in its terminal head, and repeated this in the following year.
The consequence was that the average number of rays increased rapidly, and with it the absolute maximum of the whole strain. The average came up from 21 to 34. Brighter and brighter crowns of the yellow rays improved my race, until it became difficult and very time consuming to count all the large rays of the borders. The largest numbers determined in the succeeding generations increased by leaps from 21 to 34 in the first year, and thence to 48 and 66 in the two succeeding summers. Every year I was able to save enough seed from the very best plant and to use it only for the continuance of the race. Before the selected plants were allowed to open the flowers from which the seed  was to be gathered, nearly the whole remaining culture was exterminated, excepting only some of the best examples, in order to have the required material for cross-pollination by insects. Each new generation was thereby as sharply selected as possible with regard to both parents.
All flower-heads were of course closely inspected. Not the slightest indication of real doubling was discovered, even in the summer of 1899 in the fourth generation of my selected race. But among the best the new character suddenly made its appearance. It was at the commencement of September (1899), too late to admit of the seeds ripening before winter. An inspection of the younger heads was made, which revealed three heads with some few rays in the midst of the disk on one plant, the result of the efforts of four years. Had the germ of the mutation lain hidden through all this time? Had it been present, though dormant in the original sample of seed? Or had an entirely new creation taken place during my continuous endeavors? Perhaps as their more or less immediate result? It is obviously impossible to answer these questions, before further and similar experiments shall have been performed, bringing to light other details that will enable us to reach a more definite conclusion.
 The fact that the origination of such forms is accessible to direct investigation is proven quite independently of all further considerations. The new variety came into existence at once. The leap may have been made by the ancestor of the year 1895, or by the plant of 1899, which showed the first central rays, or the sport may have been gradually built up during those four years. In each case there was a leap, contrasting with the view which claims a very long succession of years for the development of every new character.
Having discovered this first trace of doubling, it was to be expected that the new variety would be at once as pure and as rich as other double composites usually are. Some effect of the crossing with the other seed-bearing individuals might still disturb this uniformity in the following year, but another year's work would eliminate even this source of impurity.
These two years have given the expected result. The average number of the rays, which had already arisen from 13 to 34 now at once came up to 47 and 55, the last figure being the sum of 21 and 34 and therefore the probable uttermost limit to be reached before absolute doubling. The maximum numbers came as high as 100 in 1900, and reached even 200 in 1901. Such heads are as completely double as are the  brightest heads of the most beautiful double commercial varieties of composites. Even the best white camomiles (Chrysanthemum inodorum) and the gold-flowers or garden-marigolds (Calendula officinalis) do not come nearer to purity since they always have scores of little tubular florets between the rays on their disks.
Real atavists or real reversionists were seen no more after the first purification of the race. I have continued my culture and secured last summer (1903) as many and as completely doubled heads as previously. The race has at once become permanent and constant. It has of course a wide range of fluctuating variability, but the lower limit has been worked up to about 34 rays, a figure never reached by the grandiflorum parent, from which my new variety is thus sharply separated.
Unfortunately the best flowers and even the best individuals of my race are wholly barren. Selection has reached its practical limit. Seeds must be saved from less dense heads, and no way has been found of avoiding it. The ray-florets are sterile, even in the wild species, and when growing in somewhat large numbers on the disk, they conceal the fertile flowers from the visiting insects, and cause them also to be sterile. The same is the case with the best cultivated forms. Their showiest individuals are  barren, and incapable of the reproduction of the race.
This last is therefore, of necessity, always continued by means of individuals whose deviation from the mean average is the least. But in many cases the varieties are so highly differentiated that selection has become quite superfluous for practical purposes. I have already discussed the question as to the actual moment, in which the change of the grandiflorum variety into the new plenum form must be assumed to have taken place. In this respect some stress is to be laid on the fact that the improvement through selection has been gradual and continuous, though very rapid from the first moment. But with the appearance of the first stray rays within the disk, this continuity suddenly changed. All the children of this original mutated plant showed the new character, the rays within the disk, without exception. Not on all the heads, nor even on the majority of the heads on some individuals, but on some heads all gave clear proof of the possession of the new attribute. This was present in all the representatives of the new race, and had never been seen in any of their parents and grandparents. Here there was evidently a sudden leap, at least in the external form of the plants. And it seems to me to be the most simple conception,  that this visible leap directly corresponded to that inner change, which brought about the complete inheritability of the new peculiarity. It is very interesting to observe how completely my experience agrees with the results of the observations of breeders at large. No doubt a comparison is difficult, and the circumstances are not adequate to a close study.
Isolation and selection have been applied commonly only so far as was consistent with the requirements of practical horticulture, and of course a determination of the hereditary percentage was never made. The disregard of this feature made necessary a greater length of time and a larger number of generations to bring about the desired changes. Notwithstanding this, however, it has been seen that double varieties are produced suddenly. This may have occurred unexpectedly or after a few years' effort toward the end desired. Whether this sudden appearance is the consequence of a single internal differentiating step, or of the rapid succession of lesser changes, cannot yet be made out. The extreme variability of double flowers and the chance of their appearance with only slight indications of the previous petaloid alterations of a few stamens may often result in their origin being overlooked, while subsequent generations may come in for full notice.  In the greater number of cases recorded it remains doubtful whether the work said to be done to obtain a new double variety was done before the appearance of these preliminary indications or afterward.
In the first case, it would correspond with our selection of large numbers of florets in the outer rays, in the second however, with the ordinary purification of new races from hybrid mixtures.
In scientific selection-experiments such crosses are of course avoided, and the process of purification is unnecessary, even as in the Chrysanthemum culture. The first generation succeeding the original plant with disk-rays was in this respect wholly uniform and true to the new type.
In practice the work does not start from such slight indications, and is done with no other purpose in view than to produce double flowers in species in which they did not already exist. Therefore it is of the highest importance to know the methods used and the chances of success. Unfortunately the evidence is very scanty on both points.
Lindley and other writers, on horticultural theory and practice assert that a large amount of nourishment tends to produce double flowers, while a culture under normal conditions,  even if the plants are very strong and healthy, has no such effect. But even here it remains doubtful whether it applies to the period before or after the internal mutation. On the other hand success is not at all to be relied upon, nor is the work to be regarded as easy. The instances of double flowers said to be obtainable at will, are too rare in comparison with the number of cases, where the first indication of them was found accidentally.
Leaving all these doubtful points, which will have to be cleared up by further scientific investigation, the high degree of variability requires further discussion. It may be considered from three different points of view according to the limit of the deviation from the average, to the dependency on external conditions and to periodicity. It seems best to take up the last two points first.
On a visit to a nursery at Erfurt I once inspected an experiment with a new double variety of the common blue-bottle or blue corn-flower. The plants were dependent on the weather to a high degree. Bad weather increased the number of poorly filled flower-heads, while warm and sunny days were productive of beautiful double flowers. The heads that are borne by strong branches have a greater tendency to become double than those of the weaker ones,  and towards the autumn, when all those of the first group are faded away, and only a weak though large section of the heads is still flowering, the whole aspect of the variety gradually retrogrades. The same law of dependency and periodicity is prevalent everywhere. In my own cultures of the improved field-marigold I have observed it frequently. The number of the ray-florets may be considered as a direct response to nourishment, both when this is determined by external circumstances, and when it depends on the particular strength of the branch, which bears the head in question. It is a case exactly similar to that of the supernumerary carpels of the pistilloid poppy, and the deductions arrived at with that variety may be applied directly to double flowers.
This dependency upon nourishment is of high practical importance in combination with the usual effect of the doubling which makes the flowers sterile. It is a general rule that the most perfect flowers do not produce seed. At the height of the flowering period the external circumstances are the most favorable, and the flowering branches still constitute the stronger axes of the plants. Hence we may infer that sterility will prevail precisely in this period. Many varieties are known to yield only seeds from the very last flowers, as for instance some  double begonias. Others bear only seed on their weaker lateral branches, as the double camomile, or become fertile only towards the fall, as is often the case with the above quoted Erfurt variety of the blue-bottle. As far as I have been able to ascertain, such seeds are quite adequate for the reproduction and perpetuation of the double varieties, but the question whether there are differences between the seeds of the more or less double flowers of the same plants still remains open. It is very probable, from a theoretical point of view, that such differences exist, but perhaps they are so slight, as to have practically no bearing on the question.
On the ground of their wide range of variability, the double varieties must be regarded as pertaining to the group of ever-sporting forms. On one side they fluctuate in the direction towards such petalomanous flowers as are borne by the stocks and others, which we have previously discussed. Here no trace of the fertile organs is left. But this extreme is never reached by petaloid double flowers. A gap remains which, often overlooked, always exists, and which sharply separates the two types. On the other hand the alteration of the stamens gradually relapses to perfectly single flowers. Here the analogy with the pistillody of the poppies and with the "five-leaved" clover is obvious.
 This conception of the inner nature of double flowers explains the fact that the varietal mark is seldom seen to be complete throughout larger groups of individuals, providing these have not been already selected by this character. Tagetes africana is liable to produce some poorly filled specimens, and some double varieties of carnations are offered for sale with the note that the seed yields only 80% of doubles. With Chrysanthemum coronarium and blue-bottles this figure is often announced to be only about 50%. No doubt it is partly due to impurities, caused by vicinism, but it is obviously improbable that the effect of these impurities should be so large.
Some cases of partial reversion may be interpreted in the same way. Among the garden anemones, Anemone coronaria, there is a variety called the "Bride," on account of its pure white dowers. It is for sale with single and with double flowers, and these two forms are known to sport into one another, although they are multiplied in the vegetative way. Such cases are known to be of quite ordinary occurrence. Of course such sports must be considered as partial, and the same stem may bear both types of flowers. It even happens that some particular flower is partly double and partly single. Mr. Krelage, of Haarlem, had the kindness to  send me such a curious flower. One half of it was completely double, while the other half was entirely single, bearing normal and fertile stamens in the ordinary number.
The same halfway doubling is recorded to occur among composites sometimes, and from the same source I possess in my collection a head of Pyrethrum roseum, bearing on half of its disk elongated corolla tubes, and on the other half the small disk-florets of the typical species.
It is a current belief, that varieties are improved by continued culture. I have never been able to ascertain the grounds on which this conviction rests. It may be referred either to the purity of the race or to the complete development of the varietal character. In the first case it is a question of hybrid mixtures from which many young varieties must be freed before being placed on the market. But as we have already seen in a former lecture, this requires only three or four years, and afterwards the degree of purity is kept up to the point which proves to be the most suitable for practical purposes. The complete development of the varietal character is a question restricted to ever-sporting varieties, since in white flowers and other constant varieties this degree is variable in a very small and unimportant measure.  Hence the double flowers seem to afford a very good example for this discussion.
It can be decided by two facts. First by a consideration of the oldest double varieties, and secondly by that of the very youngest. Are the older ones now in a better condition than at the outset? Have they really been gradually improved during the centuries of their existence? Obviously this can only be answered by a comparison of the figures given by older writers, with the varieties as they are now in culture. Munting's drawings and descriptions are now nearly two centuries and a half old, but I do not find any real difference between his double varieties and their present representatives. So it is in other cases in which improvements by crossing or the introduction of new forms does not vitiate the evidence. Double varieties, as a rule, are exactly the same now, as they were at the time of their first introduction.
If this were otherwise one would expect that young double varieties should in the main display only slight grades of the anomaly, and that they would require centuries to reach their full development. Nothing of the kind is on record. On the contrary the newest double sorts are said to be not only equal to their predecessors, but to excel them. As a rule such claims may be exaggerated, but not to any great extent.  This is proven in the simplest way by the result of our own experiment.
In the double field-marigold we have the very first generation of a variety of pure and not hybrid origin. It shows the new attribute in its full development. It has flower-heads nearly as completely filled as the best double varieties of allied cultivated composites. In the second generation it reached heads with 200 rays each, and much larger numbers will seldom be seen in older species on heads of equal size. I have compared my novelty with the choicest double camomiles and others, but failed to discover any real difference. Improvement of the variety developed in the experiments carried on by myself seems to he excluded by the fact that it comes into conflict with the same difficulty that confronts the older cultivated species, viz.: the increasing sterility of the race.
It is perfectly evident that this double marigold is now quite constant. Continuously varying about a fixed average it may live through centuries, but the mean and the limits will always remain the same, as in the case of the ever-sporting varieties.
Throughout this lecture I have spoken of double flowers and double flower-heads of composites as of one single group. They are as nearly related from the hereditary point of  view, as they are divergent in other respects. It would be superfluous to dwell any longer upon the difference between heads and flowers. But it is as well to point out, that the term double flowers indicates a motley assemblage of different phenomena. The hen-and-chicken daisy, and the corresponding variety of the garden cineraria (Cineraria cruenta), are extremes on one side. The hen-and-chicken type occurs even in other families and is known to produce most curious anomalies, as with Scabiosa, the supernumerary heads of which may be produced on long stalks and become branched themselves in the same manner.
Petalody of the stamens is well known to be the ordinary type of doubling. But it is often accompanied by a multiplication of the organs, both of the altered stamens and of the petals themselves. This proliferation may consist in median or in lateral cleavages, and in both cases the process may be repeated one or more times. It would be quite superfluous to give more details, which may be gathered from any morphologic treatise on double flowers. But from the physiologic point of view all these cases are to be considered as one large group, complying with previously given definitions of the ever-sporting varieties. They are very variable and wholly permanent. Obviously this  permanency agrees perfectly with the conception of their sudden origin.