The inner membrane of the ovulum, however, in general appears to be of greater importance as connected with fecundation, than as affording protection to the nucleus at a more advanced period. For in many cases, before impregnation, its perforated apex projects beyond the aperture of the testa, and in some plants puts on the appearance of an obtuse, or even dilated stigma; while in the ripe seed it is often either entirely obliterated, or exists only as a thin film, which might readily be mistaken for the epidermis of a third membrane then frequently observable.
This third coat is formed by the proper membrane or cuticle of the Nucleus, from whose substance in the unimpregnated ovulum it is never, I believe, separable, and at that period is very rarely visible. In the ripe seed it is indistinguishable from the inner membrane only by its apex, which is never perforated, is generally acute and more deeply coloured, or even sphacelated.
The membrane of the nucleus usually constitutes the innermost coat of the seed. But in a few plants an additional coat, apparently originating in the inner membrane of Grew, the vesicula colliquamenti or amnios of Malpighi also exists.
In general the Amnios, after fecundation, gradually enlarges, till at length it displaces or absorbs the whole substance of the nucleus, containing in the ripe seed both the embryo and albumen, where the latter continues to exist. In such cases, however, its proper membrane is commonly obliterated, and its place supplied either by that of the nucleus, by the inner membrane of the ovulum, or, where both these are evanescent, by the testa itself.
In other cases the albumen is formed by a deposition of granular matter in the cells of the nucleus. In some of these cases the membrane of the amnios seems to be persistent, forming even in the ripe seed a proper coat for the embryo, the original attachment of whose radicle to the apex of this coat may also continue. This, at least, seems to me the most probable explanation of the structure of true Nymphaeaceae, namely, Nuphar, Nymphaea, Euryale, Hydropeltis, and Cabomba, notwithstanding their very remarkable germination, as observed and figured in Nymphaea and Nuphar by Tittmann.*
(*Footnote. Keimung der Pflanzen page 19 et 27 table 3 et 4.)
In support of this explanation, which differs from all those yet given, I may here advert to an observation published many years ago, though it seems to have escaped every author who has since written on the subject, namely, that before the maturity of the seed in Nymphaeaceae, the sacculus contains along with the embryo a (pulpy or semi-fluid) substance, which I then called Vitellus, applying at that time this name to every body interposed between the albumen and embryo.* The opinion receives some confirmation also from the existence of an extremely fine filament, hitherto overlooked, which, originating from the centre of the lower surface of the sacculus, and passing through the hollow axis of the Albumen, probably connects this coat of the Embryo in an early stage with the base of the nucleus.
(*Footnote. Prodr. Flor. Nov. Holl. 1 page 306.)
The same explanation of structure applies to the seeds of Piperaceae and Saururus; and other instances occur of the persistence either of the membrane or of the substance of the amnios in the ripe seed.
It may be concluded from the whole account which I have given of the structure of the ovulum, that the more important changes consequent to real, or even to spurious fecundation, must take place within the nucleus: and that the albumen, properly so called, may be formed either by a deposition or secretion of granular matter in the utriculi of the amnios, or in those of the nucleus itself, or lastly, that two substances having these distinct origins, and very different textures, may co-exist in the ripe seed, as is probably the case in Scitamineae.