deeply divided leaves with a much smaller surface-area (Fig. 8). Most of the common inland plants found on sand-dunes possess smaller leaves than when they grow on more hospitable ground. Nor must we forget that the semi-prostrate form so frequently adopted by dune-plants is still another method by which excessive transpiration can be reduced, since every extra inch in height exposes the plant more to the desiccating action of the strong winds.
FIG. 8.—Different leaf-forms of the red-fruited dandelion: a. Young leaf from a plant growing with abundant moisture. b. Leaf from a plant growing on dry sand.
It will be clear from what has been said that many plants are capable of modifying their normal form when growing in dry habitats. Of the various transpiration-checks which have been described, undoubtedly the development of a thick cuticle is the most frequent one employed by coastal plants. It possesses an added importance for plants inhabiting open sand-dunes in that it also protects them from possible injury caused by the sand being blown against them. Anyone who has done any botanising on exposed sand-dunes during a high wind will know how violent this bombardment can be!
It is important to point out, before we leave this subject, that it is only during periods of water-shortage that these mechanisms for reducing transpiration become important. Recent research has shown that xerophytes transpire during wet spells at least as much as, and often more than, ordinary plants. In those cases where the transpiration-rate becomes unusually high, it may be related to the necessity for rapid growth and carbon assimilation during the infrequent wet periods. What really characterises a xerophyte is that it can, if need be, decrease its transpiration-rate to a minimum when living under drought conditions. In addition, the actual protoplasm (living matter) seems able to withstand desiccation to an unusual extent.
SUCCULENCE
Some xerophytes employ quite a different method to provide against water shortage, though it is often found in combination with the leaf-modifications already described. It will be noticed that many plants growing in dry places have a fleshy or succulent appearance. This is due to the development of large colourless cells, known collectively as “aqueous tissue,” which are employed for storing water. This is usually confined to the leaves as in the stonecrops (Sedum spp.) (Pl. 14) or the sea-spurge (Euphorbia paralias) (Pl. XIX), but sometimes the whole stem is succulent as in the glassworts (Salicornia spp.) (Fig. 9(b)) or the familiar Cacti, the leaves in these cases being reduced to mere scales or spines. As a rule these cells occupy the centre of the leaf or stem, and the green cells which are used for photosynthesis occur nearer the edges. In dry weather, as water is gradually lost by transpiration or by its passage into the green cells, the water-holding cells shrink; when the water-supply improves, they expand once more. They function, in fact, as water-storage cisterns for use by the plant in times of drought. Desert succulents, which often possess extremely thick cuticles to reduce transpiration, can exist for prolonged periods without an external supply of water, during which they gradually shrivel until they can replace their internal water supplies when the rain comes.
It is rather surprising that halophytes should form the largest class of plants exhibiting succulence in this country. It has already been pointed out that the chief characteristics of this group are that they can exert sufficiently large osmotic pressures to withdraw water from a soil which is saturated with sea-water, and that the protoplasm forming their cells is not injured by exposure to salt solutions. Under normal conditions, therefore, they should not encounter much trouble with their water-supplies, and it is difficult to understand why they should develop aqueous tissue so extensively. It is possible that they draw on their internal reserves of water when the concentration of salt in the soil alters too rapidly for them to accommodate their osmotic pressure to it, but this can hardly account for such a widespread characteristic. Furthermore, it has been shown that halophytes do not, in fact, withstand drought like succulent xerophytes, but actually wither quite rapidly.
The most likely explanation is that the similarity in appearance of succulent xerophytes and halophytes is largely accidental. There is considerable evidence to suggest that the latter become succulent as a result of some chemical effect associated with salt, probably with the chlorine rather than the sodium part of it (salt is a simple compound between these two elements). All plants when growing in a saline soil absorb some salt, for the cell-walls of the root-hairs never function as “perfect” semipermeable membranes, but allow a certain amount of the dissolved substances to pass through them. This is, of course, true of all types of plant, for otherwise they would be unable to obtain the small quantities of other mineral salts essential to their growth. Many halophytes, however, absorb very large amounts of common salt; the ash of some of them, like the glassworts, was formerly used on a large scale to provide soda for glass-making, and certain plants, as for example thrift, actually excrete surplus salt from the glands on their leaves. That the development of succulent leaves is closely connected with the absorption of salt is borne out by the behaviour of many non-halophytes when they grow in places exposed to sea-water. Many inland plants found on open beaches or on cliffs within reach of sea-spray possess much more fleshy leaves than they have in their normal habitats; bird’s-foot trefoil (Lotus corniculatus), kidney vetch (Anthyllis vulneraria) and the greater knapweed (Centaurea scabiosa) are species which often show this effect. Some years ago I analysed the ash of certain inland plants which had been exposed to sea-spray in this way and found 13.5 per cent of salt in that of the kidney vetch. Evidence such as this points strongly to the conclusion that some chemical action connected with salt is the primary cause of succulence in halophytes, and there seems little reason to associate it with the problem of conserving water. The similarity between succulent xerophytes and halophytes is remarkable and we must leave it at that.
FIG. 9.—Comparison of root-systems of annual glasswort and rice-grass:-a. Small plant of annual glasswort; b. Two joints from stem of annual glasss wort enlarged to show leaf-scales; c. Base of a rice-grass plant showing vertical anchoring roots and horizontal feeding roots and stolons (Fig. c from Tansley after Oliver, 1926).
ROOT-SYSTEMS
The roots of coastal plants are very characteristic and present many points of interest. The majority of true halophytes (i.e. plants which normally grow where the soil-water is saline) possess very deep roots, generally markedly woody. Most of these plants are perennials and the chief value of their long roots in a salt-marsh is to enable them to secure a firm anchorage in relatively unstable mud. It also allows them to derive their main water supplies from regions where the concentration of salt is less variable than it is in the surface layers. Annual glasswort (Salicornia stricta) (Pl. XIII) possesses only quite short roots, and as a result is liable to become dislodged from the unstable mud if there is a strong tidal flow (Fig. 9(a)). This is in marked contrast to rice-grass (Spartina townsendii) (Pl. XIV), which occupies much the same position as a pioneer colonist in many of the south coast salt-marshes. This plant develops a most extensive root-system and becomes so firmly anchored in the mud that it can easily resist the strong currents produced by the ebb and flow of the tides (Fig. 9(c)). Its remarkable powers of spreading over soft mud and stabilising the surface are described more fully on page 71. Plants growing on sea-cliffs also develop very long roots, which serve the dual purpose of anchoring them firmly against uprooting by the violent winds encountered in these exposed places and of enabling them to tap deep-seated supplies of water. Well-established plants of thrift or samphire (Crithmum maritimum) (Pl. 10) frequently possess roots several feet long,