Here belongs Euphorbia paralias, a British littoral plant, the buoyant seeds of which occur in the stranded seed-drift of English and Mediterranean beaches.
In the following general discussion of the groups, reference will be made only to the plants best illustrating the different varieties of structure connected with buoyancy; whilst mention of the other plants will in some cases be found in other parts of this volume, as shown in the Index; and the matter is discussed at some length in not a few of the species.
The First Group.
Of the first group, where the floating power is due to the unoccupied space in the cavity of the seed or fruit, the Convolvulaceæ offer the most typical examples. Here as a rule the crumpled embryo fills the seed-cavity more or less incompletely; and it is on the relative size of the unoccupied space that the sinking or floating of the seed depends. In those plants where the seed sinks the seed-cavity may be almost filled, as in Ipomœa tuberculata, or densely packed, as in Ipomœa pentaphylla, and in species of Cuscuta. When the seed floats, as with Ipomœa pes capræ, I. glaberrima, &c., the unoccupied space is relatively large; and when, as with I. bona nox and I. turpethum, the behaviour of the seeds is irregular, some floating, and others sinking, a corresponding variation exists in the extent to which the seed-cavity is filled. This applies also to the irregular behaviour of the seeds of Ipomœa peltata and of Argyreia tiliæfolia. A singular instance is afforded by the seeds of Ipomœa insularis, collected by me in Fiji and Hawaii. Those from Fiji were incompletely filled, and consequently buoyant. Those from Hawaii were more densely packed and sank.... The three British species of Convolvulus illustrate the same principle, namely, C. arvensis, with non-buoyant seeds; C. soldanella, with buoyant seeds; and C. sepium, with seeds irregular in behaviour.
In the case of plants of the Convolvulaceæ, possessing buoyant seeds, there is always evidence of marked shrinking of the seed-contents before the final setting and hardening of the seed-coats. The embryo often appears shrivelled and dried up, and is almost brittle, so that large spaces are produced in the seed-cavity. If we partly divide such a seed and place it in water, the embryo absorbs water rapidly, and within an hour is soft, healthy-looking, and much swollen, the interspaces being filled with a jelly-like mucilage. It is therefore evident that absolute impermeability of the seed-coats is essential for the successful transport by sea-currents of the floating seed; and we can only suppose that the shrinking of the seed-contents takes place before the final setting of the tests. That with the buoyant seeds the coats are quite waterproof was illustrated in many of my experiments where, after a period of flotation covering several months, and sometimes a year or more, the seed-contents were still quite dry and shrunken. The limit of buoyancy, as I have shown in [Chapter IX.], depends on an attempt at germination on the part of the floating seed, which then absorbs water, softens, swells, and sinks.
It is, therefore, not a matter of surprise that non-buoyant seeds of the Convolvulaceæ do not gain floating power after prolonged drying of many months. It is also to be expected that, as we find in Fiji, when a characteristic shore-species with buoyant seeds like Ipomœa pes capræ extends far inland, the seeds retain their floating powers. Seed-buoyancy of this description is, on the face of it, purely mechanical.
EXPLANATION OF THE DIAGRAMS ILLUSTRATING THE CAUSES OF SEED-BUOYANCY
1. Entada scandens (natural size): (a), the shell; (b), the kernel; (c), the intercotyledonary cavity. The shell consists of three coats—an outer and an inner hard chitinous coat, and an intermediate layer of brown cellular tissue containing little or no air. The buoyancy is due entirely to the central cavity, neither the seed-tests nor the seed contents possessing any floating power (see page [181]).
2. Mucuna urens, from Hawaii (natural size). The kernel (b) sinks, and the shell has no floating power except where it possesses (under the raphe) a layer of dark brown, air-bearing, spongy tissue (a). This, however, is not sufficiently developed to endow the seed with buoyancy, which is due to the intercotyledonary cavity (c). (see page [111]).
3., 4. Mucuna gigantea, from Fiji (natural size). The kernel (b) sinks, and the seed owes its floating power entirely to the existence in the shell (a) of a layer of brown, spongy, air-bearing tissue which is mostly developed at the circumference and is almost wanting at the flat sides of the seed (see page [115]).