But Sprengel had divined but half the truth. The insect was necessary, it was true, but the Sprengel idea was concerned only with the individual flower, and the great botanist was soon perplexed and confounded by an opposing array of facts which completely destroyed the authority of his work-facts which showed conclusively that the insect could not thus convey the pollen as described, because the stigma in the flower was either not yet ready to receive it-perhaps tightly closed against it-or was past its receptive period, even decidedly withered.
[Illustration]
This radical assumption of fertilization in the individual flower, which lay at the base of Sprengel's theory, thus so completely exposed as false, discredited his entire work. The good was condemned with the bad, and the noble volume was lost in comparative oblivion-only to be finally resurrected and its full value and significance revealed by the keen scientific insight of Darwin (1859). From the new stand-point of evolution through natural selection the facts in Sprengel's work took on a most important significance. Darwin now reaffirmed the Sprengel theory so far as the necessity of the insect was concerned, but showed that all those perplexing floral conditions which had disproved Sprengel's assumption, instead of having for their object the conveying of pollen to the stigma of the same flower, implied its transfer to the stigma of another, cross-fertilization being the evident design, or evolved and perpetuated advantage.
This solution was made logical and tenable only on the assumption that such evolved conditions, insuring cross-fertilization, were of distinct advantage to the flower in the competitive struggle for existence, and that all cross-fertilized flowers were thus the final result of natural selection.
The early ancestors of this flower were self-fertilized; a chance seedling at length, among other continual variations, showed the singular variation of ripening its stigma in advance of its pollen-or other condition insuring cross-fertilization-thus acquiring a strain of fresh vigor. The seedlings of this flower, coming now into competition with the existing weaker self-fertilized forms, by the increased vigor won in the struggle of their immediate surroundings, and inheriting the peculiarity of their parent, showed flowers possessing the same cross-fertilizing device. The seeds from these, again scattering, continued the unequal struggle in a larger and larger field and in increasing numbers, continually crowding out all their less vigorous competitors of the same species, at length to become entire masters of the field and the only representatives left to perpetuate the line of descent.
Thus we find in almost every flower we meet some astonishing development by which this cross-fertilization is effected, by which the transferrence of the pollen from one flower to the stigma of another is assured, largely through the agency of insects, frequently by the wind and water, occasionally by birds. In many cases this is assured by the pollen-bearing flowers and stigmatic flowers being entirely distinct, as in cucumbers and Indian-corn; perhaps on different plants, as in the palms and willows; again by the pollen maturing and disseminating before the stigma is mature, as already mentioned, and vice versa.
From these, the simplest forms, we pass on to more and more complicated conditions, anomalies of form and structure-devices, mechanisms, that are past belief did we not observe them in actuality with our own eyes, as well as the absolutely convincing demonstration of the intention embodied: exploding flowers, shooting flowers, flower-traps, stamen embraces, pollen showers, pollen plasters, pollen necklaces, and floral pyrotechnics-all demonstrations in the floral etiquette of welcome and au revoir to insects.
From the simplest and regular types of flowers, as in the buttercup, we pass on to more and more involved and unsymmetrical forms, as the columbine, monk's-hood, larkspur, aristolochia, and thus finally to the most highly specialized or involved forms of all, as seen in the orchid-the multifarious, multiversant orchid; the beautiful orchid; the ugly orchid; the fragrant orchid; the fetid orchid; the graceful, homely, grotesque, uncanny, mimetic, and, until the year 1859, the absolutely non-committal and inexplicable flower; the blossom which had waited through the ages for Darwin, its chosen interpreter, ere she yielded her secret to humanity.
And what is an orchid? How are we to know that this blossom which we plucked is an orchid? The average reader will exclaim, "Because it is an air-plant"-the essential requisite, it would seem, in the popular mind. Of over 3000 known species of orchids, it is true a great majority are air-plants, or epiphytes-growing upon trees and other plants, obtaining their sustenance from the air, and not truly parasitic; but of the fifty-odd native species of the northeastern United States, not one is of this character, all growing in the ground, like other plants. It is only by the botanical structure of the flowers that the orchid may be readily distinguished, the epiphytic character being of little significance botanically.
A brief glance at this structural peculiarity may properly precede our more elaborate consideration of a few species of these remarkable flowers.
The orchids are usually very irregular, and six-parted. The ovary is one-celled, and becomes a pod containing an enormous yield of minute, almost spore-like, seeds (Fig. 3) in some species, as in the vanilla pod, to the number of a million, and in one species of the maxillaria, as has been carefully computed, 1,750,000.
The pollen, unlike ordinary flowers, is gathered together in waxy masses of varying consistency, variously formed and disposed in the blossom, its grains being connected with elastic cobwebby threads, which occasionally permit the entire mass to be stretched to four or five times its length, and recover its original shape when released. This is noticeable specially in the O. spectabilis, later described. The grains thus united are readily disentangled from their mass when brought into contact with a viscid object, as, for instance, the stigma.
[Illustration: Fig. 1]
But the most significant botanical contrast and distinction is found in the union of the style and stamens in one organ, called the column (Fig. 2), the stigma and the pollen being thus disposed upon a single common stalk. The contrast to the ordinary flower will be readily appreciated by comparison of the accompanying diagrams (Fig. 1).
When, therefore, we find a blossom with the anthers or pollen receptacle united to a stalk upon which the stigma is also placed, we have an orchid.
The order is further remarkable, as Darwin first demonstrated in his wonderful volume "The Fertilization of Orchids," in that the entire group, with very few exceptions, are absolutely dependent upon insects for their perpetuation through seed. They possess no possible resource for self-fertilization in the neglect of these insect sponsors.
[Illustration: Fig. 2 a. Anther. s. Stigma.]
Many of our common wild flowers, as perfectly and effectually planned for cross-fertilization as the orchids, do retain the reserve power of final self-fertilization if unfertilized by foreign pollen.
But the orchid has lost such power, and in the progress of evolution has gradually adapted itself to the insect, often to a particular species of insect, its sole sponsor, which natural selection has again gradually modified in relation to the flower.
The above work by Darwin was mostly concerned with foreign species, generally under artificial cultivation, and so startling were the disclosures concerning these hitherto sphinx-like floral beings that a most extensive bibliography soon attested the widespread inspiration and interest awakened by its pages.