the carpel of the same or another flower. The carpel, or aggregate of carpels forming the pistil or gynaeceum, comprises an ovary containing one or more ovules and a receptive surface or stigma; the stigma is sometimes carried up on a style. The mature pollen-grain is, like other spores, a single cell; except in the case of some submerged aquatic plants, it has a double wall, a thin delicate wall of unaltered cellulose, the endospore or intine, and a tough outer cuticularized exospore or extine. The exospore often bears spines or warts, or is variously sculptured, and the character of the markings is often of value for the distinction of genera or higher groups. Germination of the microspore begins before it leaves the pollen-sac. In very few cases has anything representing prothallial development been observed; generally a small cell (the antheridial or generative cell) is cut off, leaving a larger tube-cell. When placed on the stigma, under favourable circumstances, the pollen-grain puts forth a pollen-tube which grows down the tissue of the style to the ovary, and makes its way along the placenta, guided by projections or hairs, to the mouth of an ovule. The nucleus of the tube-cell has meanwhile passed into the tube, as does also the generative nucleus which divides to form two male- or sperm-cells. The male-cells are carried to their destination in the tip of the pollen-tube.
The ovary contains one or more ovules borne on a placenta, which is generally some part of the ovary-wall. The development of the ovule, which represents the macrosporangium, is very similar to the process in Gymnosperms; when mature it consists of one or twoPistil and embryo-sac. coats surrounding the central nucellus, except at the apex where an opening, the micropyle, is left. The nucellus is a cellular tissue enveloping one large cell, the embryo-sac or macrospore. The germination of the macrospore consists in the repeated division of its nucleus to form two groups of four, one group at each end of the embryo-sac. One nucleus from each group, the polar nucleus, passes to the centre of the sac, where the two fuse to form the so-called definitive nucleus. Of the three cells at the micropylar end of the sac, all naked cells (the so-called egg-apparatus), one is the egg-cell or oosphere, the other two, which may be regarded as representing abortive egg-cells (in rare cases capable of fertilization), are known as synergidae. The three cells at the opposite end are known as antipodal cells and become invested with a cell-wall. The gametophyte or prothallial generation is thus extremely reduced, consisting of but little more than the male and female sexual cells—the two sperm-cells in the pollen-tube and the egg-cell (with the synergidae) in the embryo-sac. At the period of fertilization the embryo-sac lies in close proximity Fertilization. to the opening of the micropyle, into which the pollen-tube has penetrated, the separating cell-wall becomes absorbed, and the male or sperm-cells are ejected into the embryo-sac. Guided by the synergidae one male-cell passes into the oosphere with which it fuses, the two nuclei uniting, while the other fuses with the definitive nucleus, or, as it is also called, the endosperm nucleus. This remarkable double fertilization as it has been called, although only recently discovered, has been proved to take place in widely-separated families, and both in Monocotyledons and Dicotyledons, and there is every probability that, perhaps with variations, it is the normal process in Angiosperms. After impregnation the fertilized oosphere immediately surrounds itself with a cell-wall and becomes the oospore which by a process of growth forms the embryo of the new plant. The endosperm-nucleus divides rapidly to produce a cellular tissue which fills up the interior of the rapidly-growing embryo-sac, and forms a tissue, known as endosperm, in which is stored a supply of nourishment for the use later on of the embryo. It has long been known that after fertilization of the egg has taken place, the formation of endosperm begins from the endosperm nucleus, and this had come to be regarded as the recommencement of the development of a prothallium after a pause following the reinvigorating union of the polar nuclei. This view is still maintained by those who differentiate two acts of fertilization within the embryo-sac, and regard that of the egg by the first male-cell, as the true or generative fertilization, and that of the polar nuclei by the second male gamete as a vegetative fertilization which gives a stimulus to development in correlation with the other. If, on the other hand, the endosperm is the product of an act of fertilization as definite as that giving rise to the embryo itself, we have to recognize that twin-plants are produced within the embryo-sac—one, the embryo, which becomes the angiospermous plant, the other, the endosperm, a short-lived, undifferentiated nurse to assist in the nutrition of the former, even as the subsidiary embryos in a pluri-embryonic Gymnosperm may facilitate the nutrition of the dominant one. If this is so, and the endosperm like the embryo is normally the product of a sexual act, hybridization will give a hybrid endosperm as it does a hybrid embryo, and herein (it is suggested) we may have the explanation of the phenomenon of xenia observed in the mixed endosperms of hybrid races of maize and other plants, regarding which it has only been possible hitherto to assert that they were indications of the extension of the influence of the pollen beyond the egg and its product. This would not, however, explain the formation of fruits intermediate in size and colour between those of crossed parents. The signification of the coalescence of the polar nuclei is not explained by these new facts, but it is noteworthy that the second male-cell is said to unite sometimes with the apical polar nucleus, the sister of the egg, before the union of this with the basal polar one. The idea of the endosperm as a second subsidiary plant is no new one; it was suggested long ago in explanation of the coalescence of the polar nuclei, but it was then based on the assumption that these represented male and female cells, an assumption for which there was no evidence and which was inherently improbable. The proof of a coalescence of the second male nucleus with the definitive nucleus gives the conception a more stable basis. The antipodal cells aid more or less in the process of nutrition of the developing embryo, and may undergo multiplication, though they ultimately disintegrate, as do also the synergidae. As in Gymnosperms and other groups an interesting qualitative change is associated with the process of fertilization. The number of chromosomes (see Plants: Cytology) in the nucleus of the two spores, pollen-grain and embryo-sac, is only half the number found in an ordinary vegetative nucleus; and this reduced number persists in the cells derived from them. The full number is restored in the fusion of the male and female nuclei in the process of fertilization, and remains until the formation of the cells from which the spores are derived in the new generation.
In several natural orders and genera departures from the course of development just described have been noted. In the natural order Rosaceae, the series Querciflorae, and the very anomalous genus Casuarina and others, instead of a single macrospore a more or less extensive sporogenous tissue is formed, but only one cell proceeds to the formation of a functional female cell. In Casuarina, Juglans and the order Corylaceae, the pollen-tube does not enter by means of the micropyle, but passing down the ovary wall and through the placenta, enters at the chalazal end of the ovule. Such a method of entrance is styled chalazogamic, in contrast to the porogamic or ordinary method of approach by means of the micropyle.
The result of fertilization is the development of the ovule into the seed. By the segmentation of the fertilized egg, now invested by cell-membrane, the embryo-plant arises. A varying number of transverse segment-walls transform it into a pro-embryo—a cellular row of which the cell nearestEmbryology. the micropyle becomes attached to the apex of the embryo-sac, and thus fixes the position of the developing embryo, while the terminal cell is projected into its cavity. In Dicotyledons the shoot of the embryo is wholly derived from the terminal cell of the pro-embryo, from the next cell the root arises, and the remaining ones form the suspensor. In many Monocotyledons the terminal cell forms the cotyledonary portion alone of the shoot of the embryo, its axial part and the root being derived from the adjacent cell; the cotyledon is thus a terminal structure and the apex of the primary stem a lateral one—a condition in marked contrast with that of the Dicotyledons. In some Monocotyledons,
