[Illustration: Fig. 1]
As far back as botanical history avails us these forms have been the same, each true to its particular species of flower, each with an underlying purpose which has a distinct and often simple reference to its form; and yet, incredible as it now seems to us, the botanist of the past has been content with the simple technical description of the feature, without the slightest conception of its meaning, dismissing it, perhaps, with passing comment upon its "eccentricity" or "curious shape." Indeed, prior to Darwin's time it might be said that the flower was as a voice in the wilderness. In 1735, it is true, faint premonitions of its present message began to be heard through their first though faltering interpreter, Christian Conrad Sprengel, a German botanist and school-master, who upon one occasion, while looking into the chalice of the wild geranium, received an inspiration which led him to consecrate his life thence-forth to the solution of the floral hieroglyphics. Sprengel, it may be said, was the first to exalt the flower from the mere status of a botanical specimen.
This philosophic observer was far in advance of his age, and to his long and arduous researches-a basis built upon successively by Andrew Knight, Köhlreuter, Herbert, Darwin, Lubbock, Müller, and others-we owe our present divination of the flowers.
In order to fully appreciate this present contrast, it is well to briefly trace the progress, step by step, from the consideration of the mere anatomical and physiological specimen of the earlier botanists to the conscious blossom of to-day, with its embodied hopes, aspirations, and welcome companionships.
Most of my readers are familiar with the general construction of a flower, but in order to insure such comprehension it is well, perhaps, to freshen our memory by reference to the accompanying diagram (Fig. 2) of an abstract flower, the various parts being indexed.
[Illustration: Fig. 2]
The calyx usually encloses the bud, and may be tubular, or composed of separate leaves or sepals, as in a rose. The corolla, or colored portion, may consist of several petals, as in the rose, or of a single one, as in the morning-glory. At the centre is the pistil, one or more, which forms the ultimate fruit. The pistil is divided into three parts, ovary, style, and stigma. Surrounding the pistil are the stamens, few or many, the anther at the extremity containing the powdery pollen.
Although these physiological features have been familiar to observers for thousands of years, the several functions involved were scarcely dreamed of until within a comparatively recent period.
In the writings of ancient Greeks and Romans we find suggestive references to sexes in flowers, but it was not until the close of the seventeenth century that the existence of sex was generally recognized.
[Illustration: Fig. 3]
In 1682 Nehemias Grew announced to the scientific world that it was necessary for the pollen of a flower to reach the stigma or summit of the pistil in order to insure the fruit. I have indicated his claim pictorially at A (Fig. 3), in the series of historical progression. So radical was this "theory" considered that it precipitated a lively discussion among the wiseheads, which was prolonged for fifty years, and only finally settled by Linnæus, who reaffirmed the facts declared by Grew, and verified them by such absolute proof that no further doubts could be entertained. The inference of these early authorities regarding this process of pollination is perfectly clear from their statements. The stamens in most flowers were seen to surround the pistil, "and of course the presumption was that they naturally shed the pollen upon the stigma," as illustrated at B in my series. The construction of most flowers certainly seemed designed to fulfil this end. But there were other considerations which had been ignored, and the existence of color, fragrance, honey, and insect association still continued to challenge the wisdom of the more philosophic seekers. How remarkable were some of those early speculations in regard to "honey," or, more properly, nectar! Patrick Blair, for instance, claimed that "honey absorbed the pollen," and thus fertilized the ovary. Pontidera thought that its office was to keep the ovary in a moist condition. Another botanist argued that it was "useless material thrown off in process of growth." Krunitz noted that "bee-visited meadows were most healthy," and his inference was that "honey was injurious to the flowers, and that bees were useful in carrying it off"! The great Linnæus confessed himself puzzled as to its function.
For a period of fifty years the progress of interpretation was completely arrested. The flowers remained without a champion until 1787, when Sprengel began his investigations, based upon the unsolved mysteries of color and markings of petals, fragrance, nectar, and visiting insects. The prevalent idea of the insect being a mere idle accessory to the flower found no favor with him. He chose to believe that some deep plan must lie beneath this universal association. At the inception of this conviction he chanced to observe in the flower of the wild geranium (G. sylvaticum) a fact which only an inspired vision could have detected-that the minute hairs at the base of the petal, while disclosing the nectar to insects, completely protected it from rain. Investigation showed the same conditions in many other flowers, and the inference he drew was further strengthened by the remarkable discovery of his "honey-guides" in a long list of blossoms, by which the various decorations of spots, rings, and converging veins upon the petals indicated the location of the nectar.
His labors were now concentrated on the work of interpretation, until at length his researches, covering a period of two or three years, were given to the world. In a volume bearing the following victorious title, "The Secrets of Nature in Forms and Fertilization of Flowers Discovered," he presented a vast chronicle of astonishing facts. The previous discoveries of Grew and Linnæus were right so far as they went-viz., "the pollen must reach the stigma"-but those learned authorities had missed the true secret of the process. In proof of which Sprengel showed that in a great many flowers, as I have shown at C (Fig. 3), this deposit of pollen is naturally impossible, owing to the relative position of the floral parts, and that the pollen could not reach the stigma except by artificial aid. He then announced his startling theory:
1. "Flowers are fertilized by insects."
2. Insects in approaching the nectar brush the pollen from the anthers with various hairy parts of their bodies, and in their motions convey it to the stigma.
But Sprengel's seeming victory was doomed to be turned to defeat. The true "secret" was yet unrevealed in his pages. He had given a poser to Linnæus (C), yet his own work abounded with similar strange inconsistencies, which, while being scarcely admitted by himself, or ingeniously explained, were nevertheless fatal to the full recognition of his wonderful researches. For seventy years his book lay almost unnoticed.
"Let us not underrate the value of a fact; it will one day flower in a truth." The defects in Sprengel's work were, after all, not actual defects. The error lay simply in his interpretation of his carefully noted facts. As Hermann Müller has said, "Sprengel's investigations afford an example of how even work that is rich in acute observation and happy interpretation may remain inoperative if the idea at its foundation is defective." What, then, was the flaw in Sprengel's work? Simply that he had seen but half the "secret" which he claimed to have "discovered." Starting to prove that insects fertilize the flowers, his carefully observed facts only served to demonstrate in many cases the reverse-that insects could not fertilize flowers in the manner he had declared. He was met at every hand, for instance, by floral problems such as are shown at E and F, where the pollen and the stigma in the same flower matured at different periods; and even though he recognized and admitted that the pollen must in many cases be transferred from one flower to another, he failed to divine that such was actually the common vital plan involved. It may readily be imagined that his great work precipitated an intense and prolonged controversy, and incited emulous investigation by the botanists of his time. Though a few of the more advanced of his followers, among them Andrew Knight (1799), Köhlreuter (1811), Herbert (1837), Gärtner (1844), clearly recognized the principle and foreshadowed the later theory of cross-fertilization, it was not until the inspired insight of Darwin, as voiced in his "Origin of Species," contemplated these strange facts and inconsistencies of Sprengel that their full significance and actual value were discovered and demonstrated, and his remarkable book, forgotten for seventy years, at last appreciated for its true worth. Alas for the irony of fate! Under Darwin's interpretation the very "defects" which had rendered Sprengel's work a failure now became the absolute witness of a deeper truth which Sprengel had failed to discern. One more short step and he had reached the goal. But this last step was reserved for the later seer. He took the fatal double problem of Sprengel-as shown at E and F, to express the consummation pictorially-and by the simple drawing of a line, as it were, as indicated between G and H, instantly reconciled all the previous perplexities and inconsistencies, thus demonstrating the fundamental plan involved in floral construction to be not merely "insect fertilization," the fatal postulate assumed by Sprengel, but cross -fertilization-a fact which, singularly enough, the latter's own pages proved without his suspicion.