Liliaceæ.—Gloriosa Plantii.—The stem of a half-grown plant continually moved, generally describing an irregular spire, but sometimes oval figures with the longer axes directed in different lines. It either followed the sun, or moved in an opposite course, and sometimes stood still before reversing its direction. One oval was completed in 3 hrs. 40 m.; of two horseshoe-shaped figures, one was completed in 4 hrs. 35 m. and the other in 3 hrs. The shoots, in their movements, reached points between four and five inches asunder. The young leaves, when first developed, stand up nearly vertically; but by the growth of the axis, and by the spontaneous bending down of the terminal half of the leaf, they soon become much inclined, and ultimately horizontal. The end of the leaf forms a narrow, ribbon-like, thickened projection, which at first is nearly straight, but by the time the leaf gets into an inclined position, the end bends downwards into a well-formed hook. This hook is now strong and rigid enough to catch any object, and, when caught, to anchor the plant and stop the revolving movement. Its inner surface is sensitive, but not in nearly so high a degree as that of the many before-described petioles; for a loop of string, weighing 1.64 grain, produced no effect. When the hook has caught a thin twig or even a rigid fibre, the point may be perceived in from 1 hr. to 3 hrs. to have curled a little inwards; and, under favourable circumstances, it curls round and permanently seizes an object in from 8 hrs. to 10 hrs. The hook when first formed, before the leaf has bent downwards, is but little sensitive. If it catches hold of nothing, it remains open and sensitive for a long time; ultimately the extremity spontaneously and slowly curls inwards, and makes a button-like, flat, spiral coil at the end of the leaf. One leaf was watched, and the hook remained open for thirty-three days; but during the last week the tip had curled so much inwards that only a very thin twig could have been inserted within it. As soon as the tip has curled so much inwards that the hook is converted into a ring, its sensibility is lost; but as long as it remains open some sensibility is retained.
Whilst the plant was only about six inches in height, the leaves, four or five in number, were broader than those subsequently produced; their soft and but little-attenuated tips were not sensitive, and did not form hooks; nor did the stem then revolve. At this early period of growth, the plant can support itself; its climbing powers are not required, and consequently are not developed. So again, the leaves on the summit of a full-grown flowering plant, which would not require to climb any higher, were not sensitive and could not clasp a stick. We thus see how perfect is the economy of nature.
Commelynaceæ.—Flagellaria Indica.—From dried specimens it is manifest that this plant climbs exactly like the Gloriosa. A young plant 12 inches in height, and bearing fifteen leaves, had not a single leaf as yet produced into a hook or tendril-like filament; nor did the stem revolve. Hence this plant acquires its climbing powers later in life than does the Gloriosa lily. According to Mohl (p. 41), Uvularia (Melanthaceæ) also climbs like Gloriosa.
These three last-named genera are Monocotyledons; but there is one Dicotyledon, namely Nepenthes, which is ranked by Mohl (p. 41) amongst tendril-bearers; and I hear from Dr. Hooker that most of the species climb well at Kew. This is effected by the stalk or midrib between the leaf and the pitcher coiling round any support. The twisted part becomes thicker; but I observed in Mr. Veitch’s hothouse that the stalk often takes a turn when not in contact with any object, and that this twisted part is likewise thickened. Two vigorous young plants of N. lævis and N. distillatoria, in my hothouse, whilst less than a foot in height, showed no sensitiveness in their leaves, and had no power of climbing. But when N. lævis had grown to a height of 16 inches, there were signs of these powers. The young leaves when first formed stand upright, but soon become inclined; at this period they terminate in a stalk or filament, with the pitcher at the extremity hardly at all developed. The leaves now exhibited slight spontaneous movements; and when the terminal filaments came into contact with a stick, they slowly bent round and firmly seized it. But owing to the subsequent growth of the leaf, this filament became after a time quite slack, though still remaining firmly coiled round the stick. Hence it would appear that the chief use of the coiling, at least whilst the plant is young, is to support the pitcher with its load of secreted fluid.
Summary on Leaf-climbers.—Plants belonging to eight families are known to have clasping petioles, and plants belonging to four families climb by the tips of their leaves. In all the species observed by me, with one exception, the young internodes revolve more or less regularly, in some cases as regularly as those of a twining plant. They revolve at various rates, in most cases rather rapidly. Some few can ascend by spirally twining round a support. Differently from most twiners, there is a strong tendency in the same shoot to revolve first in one and then in an opposite direction. The object gained by the revolving movement is to bring the petioles or the tips of the leaves into contact with surrounding objects; and without this aid the plant would be much less successful in climbing. With rare exceptions, the petioles are sensitive only whilst young. They are sensitive on all sides, but in different degrees in different plants; and in some species of Clematis the several parts of the same petiole differ much in sensitiveness. The hooked tips of the leaves of the Gloriosa are sensitive only on their inner or inferior surfaces. The petioles are sensitive to a touch and to excessively slight continued pressure, even from a loop of soft thread weighing only the one-sixteenth of a grain (4.05 mg.); and there is reason to believe that the rather thick and stiff petioles of Clematis flammula are sensitive to even much less weight if spread over a wide surface. The petioles always bend towards the side which is pressed or touched, at different rates in different species, sometimes within a few minutes, but generally after a much longer period. After temporary contact with any object, the petiole continues to bend for a considerable time; afterwards it slowly becomes straight again, and can then re-act. A petiole excited by an extremely slight weight sometimes bends a little, and then becomes accustomed to the stimulus, and either bends no more or becomes straight again, the weight still remaining suspended. Petioles which have clasped an object for some little time cannot recover their original position. After remaining clasped for two or three days, they generally increase much in thickness either throughout their whole diameter or on one side alone; they subsequently become stronger and more woody, sometimes to a wonderful degree; and in some cases they acquire an internal structure like that of the stem or axis.
The young internodes of the Lophospermum as well as the petioles are sensitive to a touch, and by their combined movement seize an object. The flower-peduncles of the Maurandia semperflorens revolve spontaneously and are sensitive to a touch, yet are not used for climbing. The leaves of at least two, and probably of most, of the species of Clematis, of Fumaria and Adlumia, spontaneously curve from side to side, like the internodes, and are thus better adapted to seize distant objects. The petioles of the perfect leaves of Tropæolum tricolorum, as well as the tendril-like filaments of the plants whilst young, ultimately move towards the stem or the supporting stick, which they then clasp. These petioles and filaments also show some tendency to contract spirally. The tips of the uncaught leaves of the Gloriosa, as they grow old, contract into a flat spire or helix. These several facts are interesting in relation to true tendrils.
With leaf climbers, as with twining plants, the first internodes which rise from the ground do not, at least in the cases observed by me, spontaneously revolve; nor are the petioles or tips of the first-formed leaves sensitive. In certain species of Clematis, the large size of the leaves, together with their habit of revolving, and the extreme sensitiveness of their petioles, appear to render the revolving movement of the internodes superfluous; and this latter power has consequently become much enfeebled. In certain species of Tropæolum, both the spontaneous movements of the internodes and the sensitiveness of the petioles have become much enfeebled, and in one species have been completely lost.
CHAPTER III.
Tendril-Bearers.
Nature of tendrils—Bignoniaceæ, various species of, and their different modes of climbing—Tendrils which avoid the light and creep into crevices—Development of adhesive discs—Excellent adaptations for seizing different kinds of supports.—Polemoniaceæ—Cobæa scandens much branched and hooked tendrils, their manner of action—Leguminosæ—Compositæ—Smilaceæ—Smilax aspera, its inefficient tendrils—Fumariaceæ—Corydalis claviculata, its state intermediate between that of a leaf-climber and a tendril-bearer.
By tendrils I mean filamentary organs, sensitive to contact and used exclusively for climbing. By this definition, spines, hooks and rootlets, all of which are used for climbing, are excluded. True tendrils are formed by the modification of leaves with their petioles, of flower-peduncles, branches, [84] and perhaps stipules. Mohl, who includes under the name of tendrils various organs having a similar external appearance, classes them according to their homological nature, as being modified leaves, flower-peduncles, &c. This would be an excellent scheme; but I observe that botanists are by no means unanimous on the homological nature of certain tendrils. Consequently I will describe tendril-bearing plants by natural families, following Lindley’s classification; and this will in most cases keep those of the same nature together. The species to be described belong to ten families, and will be given in the following order:—Bignoniaceæ, Polemoniaceæ, Leguminosæ, Compositæ, Smilaceæ, Fumariaceæ, Cucurbitaceæ, Vitaceæ, Sapindaceæ, Passifloraceæ. [85]