Chapter 5

In other cases the differing forms are observed to grow near each other, sometimes in neighboring provinces, sometimes in the same locality, growing and flowering in mixtures of two or three or even more elementary types. The violets exhibit widespread ancient types, from which the local species may be taken to have arisen. The common ancestors of the Whitlow-grasses are probably not to be found [64] among existing forms, but numerous types are crowded together in the southern part of central Europe and more thinly scattered elsewhere, even as far as western Asia. There can be little doubt that their common origin is to be sought in the center of their geographic distribution.

Numerous other cases exhibit smaller numbers of elementary units within a systematic species; in fact purely uniform species seem to be relatively rare. But with small numbers there are of course no indications to be expected concerning their common origin or the starting point of their distribution.

It is manifest that these experiences with wild species must find a parallel among cultivated plants. Of course cultivated plants were originally wild and must have come under the general law. Hence we may conclude that when first observed and taken up by man, they must already have consisted of sundry elementary subspecies. And we may confidently assert that some must have been rich and others poor in such types.

Granting this state of things as the only probable one, we can easily imagine what must have been the consequences. If a wild species had been taken into cultivation only once, the cultivated form would have been a single elementary [65] type. But it is not very likely that such partiality would occur often. The conception that different tribes at different times and in distant countries would have used the wild plants of their native regions seems far more natural than that all should have obtained plants for cultivation from the same source or locality. If this theory may be relied upon, the origin of many of the more widely cultivated agricultural plants must have been multiple, and the number of the original elementary species of the cultivated types must have been so much the larger, the more widely distributed and variable the plants under consideration were before the first period of cultivation.

Further it would seem only natural to explain the wide variability of many of our larger agricultural and horticultural stocks by such an incipient multiformity of the species themselves. Through commercial intercourse the various types might have become mixed so as to make it quite impossible to point out the native localities for each of them.

Unfortunately historical evidence on this point is almost wholly lacking. The differences in question could not have been appreciated at that remote period, and interest the common observer but little even today. The history of most of the cultivated plants is very obscure, [66] and even the most skillful historians, by sifting the evidence afforded by the older writers, and that obtained by comparative linguistic investigations have been able to do little more than frame the most general outline of the cultural history of the most common and most widely used plants.

Some authors assume that cultivation itself might have been the principal cause of variability, but it is not proved, nor even probable, that cultivated plants are intrinsically more variable than their wild prototypes. Appearances in this case are very deceptive. Of course widely distributed plants are as a rule richer in subspecies than forms with limited distribution, and the former must have had a better chance to be taken into cultivation than the latter. In many cases, especially with the more recent cultivated species, man has deliberately chosen variable forms, because of their greater promise. Thirdly, wide variability is the most efficient means of acclimatization, and only species with many elementary units would have offered the adequate material for introduction into new countries.

From this discussion it would seem that it is more reasonable to assert that variability is one of the causes of the success of cultivation, than to assume that cultivation is a cause of variability [67] at large. And this assumption would be equally sufficient to explain the existing conditions among cultivated plants.

Of course I do not pretend to say that cultivated plants should be expected to be less variable than in the wild state, or that swarms of elementary species might not be produced during cultivation quite as well as before. However the chance of such an event, as is easily seen, cannot be very great, and we shall have to be content with a few examples of which the coconut is a notable one.

Leaving this general discussion of the subject, we may take up the example of the beets. The sugar-beet is only one type from among a horde of others, and though the origin of all the single types is not historically known, the plant is frequently found in the wild state even at the present time, and the native types may be compared with the corresponding cultivated varieties.

The cultivation of beets for sugar is not of very ancient date. The Romans knew the beets and used them as vegetables, both the roots and the leaves. They distinguished a variety with white and one with red flesh, but whether they cultivated them, or only collected them from where they grew spontaneously, appears to be unknown.

[68] Beets are even now found in large quantities along the shores of Italy. They prefer the vicinity of the sea, as do so many other members of the beet family, and are not limited to Italy, but are found growing elsewhere on the littoral of the Mediterranean, in the Canary Islands and through Persia and Babylonia to India. In most of their native localities they occur in great abundance.

The color of the foliage and the size of the roots are extremely variable. Some have red leafstalks and veins, others a uniform red or green foliage, some have red or white or yellow roots, or exhibit alternating rings of a red and of a white tinge on cut surfaces. It seems only natural to consider the white and the red, and even the variegated types as distinct varieties, which in nature do not transgress their limits nor change into one another. In a subsequent lecture I will show that this at least is the rule with the corresponding color-varieties in other genera.

The fleshiness or pulpiness of the roots is still more variable. Some are as thick as the arm and edible, others are not thicker than a finger and of a woody composition, and the structure of this woody variety is very interesting. The sugar-beet consists, as is generally known, of concentric layers of sugar-tissue and of vascular [69] strands; the larger the first and the smaller the latter, the greater is, as a rule, the average amount of sugar of the race. Through the kindness of the late Mr. Rimpau, a well known German breeder of sugar-beet varieties, I obtained specimens from seed of a native wild locality near Bukharest. The plants produced quite woody roots, showing almost no sugar tissue at all. Woody layers of strongly developed fibrovascular strands were seen to be separated one from another only by very thin layers of parenchymatous cells. Even the number of layers is variable; it was observed to be five in my plants; but in larger roots double this number and even more may easily be met with.

Some authors have distinguished specific types among these wild forms. While the cultivated beets are collected under the head of _Beta vulgaris_, separate types with more or less woody roots have been described as _Beta maritima_ and _Beta patula_. These show differences in the habit of the stems and the foliage. Some have a strong tendency to become annual, others to become biennial. The first of course do not store a large quantity of food in their roots, and remain thin, even at the time of flowering. The biennial types occur in all sizes of roots. In the annuals the stems may vary from [70] erect to ascending, and the name _patula_ indicates stems which are densely branching from the base with widely spreading branches throughout. Mr. Em. von Proskowetz of Kwassitz, Austria, kindly sent me seeds of this _Beta patula_, the variability of which was so great in my cultures as to range from nearly typical sugar-beets to the thin woody type of Bukharest.

Broad and narrow leaves are considered to be differentiating marks between _Beta vulgaris_ and _Beta patula_, but even here a wide range of forms seem to occur.

Rimpau, Proskowetz, Schindler and others have made cultures of beets from wild localities in order to discover a hypothetical common ancestor of all the present cultivated types. These researches point to the _B. patula_ as the probable ancestor, but of course they were not made to decide the question as to whether the origination of the several now existing types had taken place before or during culture. From a general point of view the variability of the wild species is parallel to that of the cultivated forms to such a degree as to suggest the multiple origin of the former. But a close investigation of this highly important problem has still to be made.

The varieties of the cultivated beets are commonly [71] included in four subspecies. The two smallest are the salad-beets and the ornamental forms, the first being used as food, and ordinarily cultivated in red varieties, the second being used as ornamental plants during the fall, when they fill the beds left empty by summer flowers, with a bright foliage that is exceedingly rich in form and color. Of the remaining subspecies, one comprises the numerous sorts cultivated as forage-crops and the other the true sugar-beets. Both of them vary widely as to the shape and the size of the roots, the quality of the tissue, the foliage and other characteristics.

Some of these forms, no doubt, have originated during culture. Most of them have been improved by selection, and no beet found in the wild state ever rivals any cultivated variety. But the improvement chiefly affects the size, the amount of sugar and nutrient substances and some other qualities which recur in most of the varieties. The varietal attributes themselves however, are more or less of a specific nature, and have no relation to the real industrial value of the race. The short-rooted and the horn-shaped varieties might best be cited as examples.

The assertion that the sundry varieties of forage-beets are not the result of artificial selection, [72] is supported in a large measure by the historic fact that the most of them are far older than the method of conscious selection of plants itself. This method is due to Louis Vilmorin and dates from the middle of the last century. But in the sixteenth century most of our present varieties of beets were already in cultivation. Caspar Bauhin gives a list of the beets of his time and it is not difficult to recognize in it a large series of subspecies and varieties and even of special forms, which are still cultivated. A more complete list was published towards the close of the same century by Olivier de Serres in his world-renowned "Theatre d'Agriculture" (Paris, 1600).

The red forage-beets which are now cultivated on so large a scale, had been introduced from Italy into France only a short time before.

From this historic evidence, the period during which the beets were cultivated from the time of the Romans or perhaps much later, up to the time of Bauhin and De Serres, would seem far too short for the production by the unguided selection of man of all the now existing types. On the other hand, the parallelism between the characters of some wild and some cultivated varieties goes to make it very probable that other varieties have been found in the same way, some in this country and others in that, [73] and have been taken into cultivation separately. Afterwards of course all must have been improved in the direction required by the needs of man.

Quite the same conclusion is afforded by apples. The facts are to some extent of another character, and the rule of the derivation of the present cultivated varieties from original wild forms can be illustrated in this case in a more direct way. Of course we must limit ourselves to the varieties of pure ancestry and leave aside all those which are of hybrid or presumably hybrid origin.

Before considering their present state of culture, something must be, said about the earlier history and the wild state of the apples.

The apple-tree is a common shrub in woods throughout all parts of Europe, with the only exception of the extreme north. Its distribution extends to Anatolia, the Caucasus and Ghilan in Persia. It is found in nearly all forests of any extent and often in relatively large numbers of individuals. It exhibits varietal characters, which have led to the recognition of several spontaneous forms, especially in France and in Germany.

The differentiating qualities relate to the shape and indumentum of the leaves. Nothing is known botanically as to differences between [74] the fruits of these varieties, but as a matter of fact the wild apples of different countries are not at all the same.

Alphonse De Candolle, who made a profound study of the probable origin of most of our cultivated plants, comes to the conclusion that the apple tree must have had this wide distribution in prehistoric times, and that its cultivation began in ancient times everywhere.

This very important conclusion by so high an authority throws considerable light on the relation between cultivated and wild varieties at large. If the historic facts go to prove a multiple origin for the cultivation of some of the more important useful plants, the probability that different varieties or elementary species have been the starting points for different lines of culture, evidently becomes stronger.

Unfortunately, this historic evidence is scanty. The most interesting facts are those concerning the use of apples by the Romans and by their contemporaries of the Swiss and middle European lake-dwellings. Oswald Heer has collected large numbers of the relics of this prehistoric period. Apples were found in large quantities, ordinarily cut into halves and with the signs of having been dried. Heer distinguished two varieties, one with large and one with small fruits. The first about 3 and [75] the other about 1.5-2 cm. in diameter. Both are therefore very small compared with our present ordinary varieties, but of the same general size as the wild forms of the present day. Like these, they must have been of a more woody and less fleshy tissue. They would scarcely have been tasteful to us, but in ancient times no better varieties were known and therefore no comparison was possible.

There is no evidence concerning the question, as to whether during the periods mentioned apples were cultivated or only collected in the wild state. The very large numbers which are found, have induced some writers to believe in their culture, but then there is no reason why they should not have been collected in quantity from wild shrubs. The main fact is that the apple was not a uniform species in prehistoric times but showed even then at least some amount of variability.

At the present day the wild apples are very rich in elementary species. Those of Versailles are not the same as those of Belgium, and still others are growing in England and in Germany. The botanical differences derived from the blossoms and the leaves are slight, but the flavor, size and shape of the fruits diverge widely. Two opinions have been advanced to explain this high degree of variability, but [76] neither of them conveys a real explanation; their aim is chiefly to support different views as to the causes of variability, and the origin of elementary species at large.

One opinion, advocated by De Candolle, Darwin and others, claims that the varieties owe their origin to the direct influence of cultivation, and that the corresponding forms found in the wild state, are not at all original, but have escaped from cultivation and apparently become wild. Of course this possibility cannot be denied, at least in any single instance, but it seems too sweeping an assertion to make for the whole range of observed forms.

The alternative theory is that of van Mons, the Belgian originator of commercial varieties of apples, who has published his experiments in a large work called "Arbres fruitiers ou Pomonomie belge." Most of the more remarkable apples of the first half of the last century were produced by van Mons, but his greatest merit is not the direct production of a number of good varieties, but the foundation of the method, by which new varieties may be obtained and improved.

According to van Mons, the production of a new variety consists chiefly of two parts. The first is the discovery of a subspecies with new desirable qualities. The second is the transformation [77] of the original small and woody apple into a large, fleshy and palatable variety. Subspecies, or what we now call elementary species were not produced by man; nature alone creates new forms, as van Mons has it. He examined with great care the wild apples of his country, and especially those of the Ardennes, and found among them a number of species with different flavors. For the flavor is the one great point, which must be found ready in nature and which may be improved, but can never be created by artificial selection. The numerous differences in flavor are quite original; all of them may be found in the wild state and most of them even in so limited a region as the Ardennes Mountains. Of course van Mons preferred not to start from the wild types themselves, when the same flavor could be met with in some cultivated variety. His general method was, to search for a new flavor and to try to bring the bearer of it up to the desired standard of size and edibility.

The latter improvement, though it always makes the impression of an achievement, is only the last stone to be added to the building up of the commercial value of the variety. Without it, the best flavored apple remains a crab; with it, it becomes a conquest. According to the method of van Mons it may be reached within [78] two or three generations, and a man's life is wholly sufficient to produce in this way many new types of the very best sorts, as van Mons himself has done. It is done in the usual way, sowing on a large scale and selecting the best, which are in their turn brought to an early maturation of their fruit by grafting, because thereby the life from seed to seed may be reduced to a few years.

Form, taste, color, flavor and other valuable marks of new varieties are the products of nature, says van Mons, only texture, fleshiness and size are added by man. And this is done in each new variety by the same method and according to the same laws. The richness of the cultivated apples of the present day was already present in the large range of original wild elementary species, though unobserved and requiring improvement.

An interesting proof of this principle is afforded by the experience of Mr. Peter M. Gideon, as related by Bailey. Gideon sowed large quantities of apple-seeds, and one seed produced a new and valuable variety called by him the "Wealthy" apple. He first planted a bushel of apple-seeds, and then every year, for nine years, planted enough seeds to produce a thousand trees. At the end of ten years all seedlings had perished except one hardy seedling [79] crab. This experiment was made in Minnesota, and failed wholly. Then he bought a small lot of seeds of apples and crab-apples in Maine and from these the "Wealthy" came. There were only about fifty seeds in the lot of crab-apple seed which produced the "Wealthy," but before this variety was obtained, more than a bushel of seed had been sown. Chance afforded a species with an unknown taste; but the growing of many thousands of seedlings of known varieties was not the best means to get something really new.

Pears are more difficult to improve than apples. They often require six or more generations to be brought from the wild woody state to the ordinary edible condition. But the varieties each seem to have a separate origin, as with apples, and the wide range of form and of taste must have been present in the wild state, long before cultivation. Only recently has the improvement of cherries, plums, currants and gooseberries been undertaken with success by Mr. Burbank, and the difference between the wild and cultivated forms has hitherto been very small. All indications point to the existence, before the era of cultivation, of larger or smaller numbers of elementary species.

The same holds good with many of the larger forage crops and other plants of great industrial [80] value. Clover exhibits many varieties, which have been cultivated indiscriminately, and often in motley mixtures. The flower heads may be red or white, large or small, cylindric or rounded, the leaves are broader or narrower, with or without white spots of a curious pattern. They may be more or less hairy and so forth. Even the seeds exhibit differences in size, shape or color, and of late Martinet has shown, that by the simple means of picking out seeds of the same pattern, pure strains of clover may be obtained, which are of varying cultural value. In this way the best subspecies or varieties may be sought out for separate cultivation. Even the white spots on the leaflets have proved to be constant characters corresponding with noticeable differences in yield.

Flax is another instance. It was already cultivated, or at least made use of during the period of the lake-dwellers, but at that time it was a species referred to as _Linum angustifolium_, and not the _Linum usitatissimum_, which is our present day flax. There are now many subspecies, elementary species, and varieties under cultivation. The oldest of them is known as the "springing flax," in opposition to the ordinary "threshing flax." It has capsules which open of themselves, in order to disseminate the seeds, while the ordinary heads of the [81] flax remain closed until the seeds are liberated by threshing. It seems probable that the first form or _Linum crepitans_ might thrive in the wild state as well as any other plant, while in the common species those qualities are lacking which are required for a normal dissemination of the seeds. White or blue flowers, high or dwarf stems, more or less branching at the base and sundry other qualities distinguish the varieties, aside from the special industrial difference of the fibres. Even the life-history varies from annual and biennial, to perennial.

It would take us too long to consider other instances. It is well known that corn, though considered as a single botanical species, is represented by different subspecies and varieties in nearly every region in which it is grown. Of course its history is unknown and it is impossible to decide whether all the tall and dwarf forms, or starchy and sweet varieties, dented or rounded kernels, and hundreds of others are older than culture or have come into existence during historic times, or as some assume, through the agency of man. But our main point now is not the origin, but only the existence of constant and sharply differentiated forms within botanical species. Nearly every cultivated plant affords instances of such diversity. Some include a few types only, while [82] others show, a large number of forms clearly separated to a greater or lesser degree.