This plant at first perplexed me much, for I continually observed its long and flexible shoots, evidently well fitted for twining, make a whole, or half, or quarter circle in one direction and then in an opposite direction; consequently, when I placed the shoots near thin or thick sticks, or perpendicularly stretched string, they seemed as if constantly trying to ascend, but always failed. I then surrounded the plant with a ma.s.s of branched twigs; the shoots ascended, and pa.s.sed through them, but several came out laterally, and their depending extremities seldom turned upwards as is usual with twining plants. Finally, I surrounded a second plant with many thin upright sticks, and placed it near the first one with twigs; and now both had got what they liked, for they twined up the parallel sticks, sometimes winding round one and sometimes round several; and the shoots travelled laterally from one to the other pot; but as the plants grew older, some of the shoots twined regularly up thin upright sticks. Though the revolving movement was sometimes in one direction and sometimes in the other, the twining was invariably from left to right; {18} so that the more potent or persistent movement of revolution must have been in opposition to the course of the sun. It would appear that this Hibbertia is adapted both to ascend by twining, and to ramble laterally through the thick Australian scrub.
I have described the above case in some detail, because, as far as I have seen, it is rare to find any special adaptations with twining plants, in which respect they differ much from the more highly organized tendril-bearers. The Solanum dulcamara, as we shall presently see, can twine only round stems which are both thin and flexible. Most twining plants are adapted to ascend supports of moderate though of different thicknesses. Our English twiners, as far as I have seen, never twine round trees, excepting the honeysuckle (Lonicera periclymenum), which I have observed twining up a young beech-tree nearly 4.5 inches in diameter. Mohl (p. 134) found that the Phaseolus multiflorus and Ipomoea purpurea could not, when placed in a room with the light entering on one side, twine round sticks between 3 and 4 inches in diameter; for this interfered, in a manner presently to be explained, with the revolving movement.
In the open air, however, the Phaseolus twined round a support of the above thickness, but failed in twining round one 9 inches in diameter. Nevertheless, some twiners of the warmer temperate regions can manage this latter degree of thickness; for I hear from Dr.
Hooker that at Kew the Ruscus androgynus has ascended a column 9 inches in diameter; and although a Wistaria grown by me in a small pot tried in vain for weeks to get round a post between 5 and 6 inches in thickness, yet at Kew a plant ascended a trunk above 6 inches in diameter. The tropical twiners, on the other hand, can ascend thicker trees; I hear from Drs. Thomson and Hooker that this is the case with the Butea parviflora, one of the Menispermaceae, and with some Dalbergias and other Leguminosae. {19} This power would be necessary for any species which had to ascend by twining the large trees of a tropical forest; otherwise they would hardly ever be able to reach the light. In our temperate countries it would be injurious to the twining plants which die down every year if they were enabled to twine round trunks of trees, for they could not grow tall enough in a single season to reach the summit and gain the light.
By what means certain twining plants are adapted to ascend only thin stems, whilst others can twine round thicker ones, I do not know. It appeared to me probable that twining plants with very long revolving shoots would be able to ascend thick supports; accordingly I placed Ceropegia Gardnerii near a post 6 inches in diameter, but the shoots entirely failed to wind round it; their great length and power of movement merely aid them in finding a distant stem round which to twine. The Sphaerostemma marmoratum is a vigorous tropical twiner; and as it is a very slow revolver, I thought that this latter circ.u.mstance might help it in ascending a thick support; but though it was able to wind round a 6-inch post, it could do this only on the same level or plane, and did not form a spire and thus ascend.
As ferns differ so much in structure from phanerogamic plants, it may be worth while here to show that twining ferns do not differ in their habits from other twining plants. In LyG.o.dium articulatum the two internodes of the stem (properly the rachis) which are first formed above the root-stock do not move; the third from the ground revolves, but at first very slowly. This species is a slow revolver: but L.
scandens made five revolutions, each at the average rate of 5 hrs. 45 m.; and this represents fairly well the usual rate, taking quick and slow movers, amongst phanerogamic plants. The rate was accelerated by increased temperature. At each stage of growth only the two upper internodes revolved. A line painted along the convex surface of a revolving internode becomes first lateral, then concave, then lateral and ultimately again convex. Neither the internodes nor the petioles are irritable when rubbed. The movement is in the usual direction, namely, in opposition to the course of the sun; and when the stem twines round a thin stick, it becomes twisted on its own axis in the same direction. After the young internodes have twined round a stick, their continued growth causes them to slip a little upwards.
If the stick be soon removed, they straighten themselves, and recommence revolving. The extremities of the depending shoots turn upwards, and twine on themselves. In all these respects we have complete ident.i.ty with twining phanerogamic plants; and the above enumeration may serve as a summary of the leading characteristics of all twining plants.
The power of revolving depends on the general health and vigour of the plant, as has been laboriously shown by Palm. But the movement of each separate internode is so independent of the others, that cutting off an upper one does not affect the revolutions of a lower one. When, however, Dutrochet cut off two whole shoots of the Hop, and placed them in water, the movement was greatly r.e.t.a.r.ded; for one revolved in 20 hrs. and the other in 23 hrs., whereas they ought to have revolved in between 2 hrs. and 2 hrs. 30 m. Shoots of the Kidney-bean, cut off and placed in water, were similarly r.e.t.a.r.ded, but in a less degree. I have repeatedly observed that carrying a plant from the greenhouse to my room, or from one part to another of the greenhouse, always stopped the movement for a time; hence I conclude that plants in a state of nature and growing in exposed situations, would not make their revolutions during very stormy weather. A decrease in temperature always caused a considerable r.e.t.a.r.dation in the rate of revolution; but Dutrochet (tom. xvii. pp.
994, 996) has given such precise observations on this head with respect to the common pea that I need say nothing more. When twining plants are placed near a window in a room, the light in some cases has a remarkable power (as was likewise observed by Dutrochet, p.
998, with the pea) on the revolving movement, but this differs in degree with different plants; thus Ipomoea jucunda made a complete circle in 5 hrs. 30 m.; the semicircle from the light taking 4 hrs.
80 m., and that towards the light only 1 hr. Lonicera brachypoda revolved, in a reversed direction to the Ipomoea, in 8 hrs.; the semicircle from the light taking 5 hrs. 23 m., and that to the light only 2 hrs. 37 m. From the rate of revolution in all the plants observed by me, being nearly the same during the night and the day, I infer that the action of the light is confined to r.e.t.a.r.ding one semicircle and accelerating the other, so as not to modify greatly the rate of the whole revolution. This action of the light is remarkable, when we reflect how little the leaves are developed on the young and thin revolving internodes. It is all the more remarkable, as botanists believe (Mohl, p. 119) that twining plants are but little sensitive to the action of light.
I will conclude my account of twining plants by giving a few miscellaneous and curious cases. With most twining plants all the branches, however many there may be, go on revolving together; but, according to Mohl (p. 4), only the lateral branches of Tamus elephantipes twine, and not the main stem. On the other hand, with a climbing species of Asparagus, the leading shoot alone, and not the branches, revolved and twined; but it should be stated that the plant was not growing vigorously. My plants of Combretum argenteum and C.
purpureum made numerous short healthy shoots; but they showed no signs of revolving, and I could not conceive how these plants could be climbers; but at last C. argenteum put forth from the lower part of one of its main branches a thin shoot, 5 or 6 feet in length, differing greatly in appearance from the previous shoots, owing to its leaves being little developed, and this shoot revolved vigorously and twined. So that this plant produces shoots of two kinds. With Periploca Graeca (Palm, p. 43) the uppermost shoots alone twine.
Polygonum convolvulus twines only during the middle of the summer (Palm, p. 43, 94); and plants growing vigorously in the autumn show no inclination to climb. The majority of Asclepiadaceae are twiners; but Asclepias nigra only "in fertiliori solo incipit scandere subvolubili caule" (Willdenow, quoted and confirmed by Palm, p. 41).
Asclepias vincetoxic.u.m does not regularly twine, but occasionally does so (Palm, p. 42; Mohl, p. 112) when growing under certain conditions. So it is with two species of Ceropegia, as I hear from Prof. Harvey, for these plants in their native dry South African home generally grow erect, from 6 inches to 2 feet in height,--a very few taller specimens showing some inclination to curve; but when cultivated near Dublin, they regularly twined up sticks 5 or 6 feet in height. Most Convolvulaceae are excellent twiners; but in South Africa Ipomoea argyraeoides almost always grows erect and compact, from about 12 to 18 inches in height, one specimen alone in Prof.
Harvey"s collection showing an evident disposition to twine. On the other hand, seedlings raised near Dublin twined up sticks above 8 feet in height. These facts are remarkable; for there can hardly be a doubt that in the dryer provinces of South Africa these plants have propagated themselves for thousands of generations in an erect condition; and yet they have retained during this whole period the innate power of spontaneously revolving and twining, whenever their shoots become elongated under proper conditions of life. Most of the species of Phaseolus are twiners; but certain varieties of the P.
multiflorus produce (Leon, p. 681) two kinds of shoots, some upright and thick, and others thin and twining. I have seen striking instances of this curious case of variability in "Fulmer"s dwarf forcing-bean," which occasionally produced a single long twining shoot.
Solanum dulcamara is one of the feeblest and poorest of twiners: it may often be seen growing as an upright bush, and when growing in the midst of a thicket merely scrambles up between the branches without twining; but when, according to Dutrochet (tom. xix. p. 299), it grows near a thin and flexible support, such as the stem of a nettle, it twines round it. I placed sticks round several plants, and vertically stretched strings close to others, and the strings alone were ascended by twining. The stem twines indifferently to the right or left. Some others species of Solanum, and of another genus, viz.
Habrothamnus, belonging to the same family, are described in horticultural works as twining plants, but they seem to possess this faculty in a very feeble degree. We may suspect that the species of these two genera have as yet only partially acquired the habit of twining. On the other hand with Tecoma radicans, a member of a family abounding with twiners and tendril-bearers, but which climbs, like the ivy, by the aid of rootlets, we may suspect that a former habit of twining has been lost, for the stem exhibited slight irregular movements which could hardly be accounted for by changes in the action of the light. There is no difficulty in understanding how a spirally twining plant could graduate into a simple root-climber; for the young internodes of Bignonia Tweedyana and of Hoya carnosa revolve and twine, but likewise emit rootlets which adhere to any fitting surface, so that the loss of twining would be no great disadvantage and in some respects an advantage to these species, as they would then ascend their supports in a more direct line. {20}
CHAPTER II.--LEAF-CLIMBERS.
Plants which climb by the aid of spontaneously revolving and sensitive petioles--Clematis--Tropaeolum--Maurandia, flower-peduncles moving spontaneously and sensitive to a touch--Rhodochiton-- Lophospermum--internodes sensitive--Solanum, thickening of the clasped petioles--Fumaria--Adlumia--Plants which climb by the aid of their produced midribs--Gloriosa--Flagellaria--Nepenthes--Summary on leaf-climbers.
We now come to our second cla.s.s of climbing plants, namely, those which ascend by the aid of irritable or sensitive organs. For convenience" sake the plants in this cla.s.s have been grouped under two sub-divisions, namely, leaf-climbers, or those which retain their leaves in a functional condition, and tendril-bearers. But these sub-divisions graduate into each other, as we shall see under Corydalis and the Gloriosa lily.
It has long been observed that several plants climb by the aid of their leaves, either by their petioles (foot-stalks) or by their produced midribs; but beyond this simple fact they have not been described. Palm and Mohl cla.s.s these plants with those which bear tendrils; but as a leaf is generally a defined object, the present cla.s.sification, though artificial, has at least some advantages.
Leaf-climbers are, moreover, intermediate in many respects between twiners and tendril-bearers. Eight species of Clematis and seven of Tropaeolum were observed, in order to see what amount of difference in the manner of climbing existed within the same genus; and the differences are considerable.
CLEMATIS.--C. glandulosa.--The thin upper internodes revolve, moving against the course of the sun, precisely like those of a true twiner, at an average rate, judging from three revolutions, of 3 hrs. 48 m.
The leading shoot immediately twined round a stick placed near it; but, after making an open spire of only one turn and a half, it ascended for a short s.p.a.ce straight, and then reversed its course and wound two turns in an opposite direction. This was rendered possible by the straight piece between the opposed spires having become rigid.
The simple, broad, ovate leaves of this tropical species, with their short thick petioles, seem but ill-fitted for any movement; and whilst twining up a vertical stick, no use is made of them.
Nevertheless, if the footstalk of a young leaf be rubbed with a thin twig a few times on any side, it will in the course of a few hours bend to that side; afterwards becoming straight again. The under side seemed to be the most sensitive; but the sensitiveness or irritability is slight compared to that which we shall meet with in some of the following species; thus, a loop of string, weighing 1.64 grain (106.2 mg.) and hanging for some days on a young footstalk, produced a scarcely perceptible effect. A sketch is here given of two young leaves which had naturally caught hold of two thin branches. A forked twig placed so as to press lightly on the under side of a young footstalk caused it, in 12 hrs., to bend greatly, and ultimately to such an extent that the leaf pa.s.sed to the opposite side of the stem; the forked stick having been removed, the leaf slowly recovered its former position.
The young leaves spontaneously and gradually change their position: when first developed the petioles are upturned and parallel to the stem; they then slowly bend downwards, remaining for a short time at right angles to the stem, and then become so much arched downwards that the blade of the leaf points to the ground with its tip curled inwards, so that the whole petiole and leaf together form a hook.
They are thus enabled to catch hold of any twig with which they may be brought into contact by the revolving movement of the internodes.
If this does not happen, they retain their hooked shape for a considerable time, and then bending upwards rea.s.sume their original upturned position, which is preserved ever afterwards. The petioles which have clasped any object soon become much thickened and strengthened, as may be seen in the drawing.
Clematis montana.--The long, thin petioles of the leaves, whilst young, are sensitive, and when lightly rubbed bend to the rubbed side, subsequently becoming straight. They are far more sensitive than the petioles of C. glandulosa; for a loop of thread weighing a quarter of a grain (16.2 mg.) caused them to bend; a loop weighing only one-eighth of a grain (8.1 mg.) sometimes acted and sometimes did not act. The sensitiveness extends from the blade of the leaf to the stem. I may here state that I ascertained in all cases the weights of the string and thread used by carefully weighing 50 inches in a chemical balance, and then cutting off measured lengths. The main petiole carries three leaflets; but their short, sub-petioles are not sensitive. A young, inclined shoot (the plant being in the greenhouse) made a large circle opposed to the course of the sun in 4 hrs. 20 m., but the next day, being very cold, the time was 5 hrs. 10 m. A stick placed near a revolving stem was soon struck by the petioles which stand out at right angles, and the revolving movement was thus arrested. The petioles then began, being excited by the contact, to slowly wind round the stick. When the stick was thin, a petiole sometimes wound twice round it. The opposite leaf was in no way affected. The att.i.tude a.s.sumed by the stem after the petiole had clasped the stick, was that of a man standing by a column, who throws his arm horizontally round it. With respect to the stem"s power of twining, some remarks will be made under C. calycina.
Clematis Sieboldi.--A shoot made three revolutions against the sun at an average rate of 3 hrs. 11 m. The power of twining is like that of the last species. Its leaves are nearly similar in structure and in function, excepting that the sub-petioles of the lateral and terminal leaflets are sensitive. A loop of thread, weighing one-eighth of a grain, acted on the main petiole, but not until two or three days had elapsed. The leaves have the remarkable habit of spontaneously revolving, generally in vertical ellipses, in the same manner, but in a less degree, as will be described under C. microphylla.
Clematis calycina.--The young shoots are thin and flexible: one revolved, describing a broad oval, in 5 hrs. 30 m., and another in 6 hrs. 12 m. They followed the course of the sun; but the course, if observed long enough, would probably be found to vary in this species, as well as in all the others of the genus. It is a rather better twiner than the two last species: the stem sometimes made two spiral turns round a thin stick, if free from twigs; it then ran straight up for a s.p.a.ce, and reversing its course took one or two turns in an opposite direction. This reversal of the spire occurred in all the foregoing species. The leaves are so small compared with those of most of the other species, that the petioles at first seem ill-adapted for clasping. Nevertheless, the main service of the revolving movement is to bring them into contact with surrounding objects, which are slowly but securely seized. The young petioles, which alone are sensitive, have their ends bowed a little downwards, so as to be in a slight degree hooked; ultimately the whole leaf, if it catches nothing, becomes level. I gently rubbed with a thin twig the lower surfaces of two young petioles; and in 2 hrs. 30 m. they were slightly curved downwards; in 5 hrs., after being rubbed, the end of one was bent completely back, parallel to the basal portion; in 4 hrs. subsequently it became nearly straight again. To show how sensitive the young petioles are, I may mention that I just touched the under sides of two with a little water-colour, which when dry formed an excessively thin and minute crust; but this sufficed in 24 hrs. to cause both to bend downwards. Whilst the plant is young, each leaf consists of three divided leaflets, which barely have distinct petioles, and these are not sensitive; but when the plant is well grown, the petioles of the two lateral and terminal leaflets are of considerable length, and become sensitive so as to be capable of clasping an object in any direction.
When a petiole has clasped a twig, it undergoes some remarkable changes, which may be observed with the other species, but in a less strongly marked manner, and will here be described once for all. The clasped petiole in the course of two or three days swells greatly, and ultimately becomes nearly twice as thick as the opposite one which has clasped nothing. When thin transverse slices of the two are placed under the microscope their difference is conspicuous: the side of the petiole which has been in contact with the support, is formed of a layer of colourless cells with their longer axes directed from the centre, and these are very much larger than the corresponding cells in the opposite or unchanged petiole; the central cells, also, are in some degree enlarged, and the whole is much indurated. The exterior surface generally becomes bright red. But a far greater change takes place in the nature of the tissues than that which is visible: the petiole of the unclasped leaf is flexible and can be snapped easily, whereas the clasped one acquires an extraordinary degree of toughness and rigidity, so that considerable force is required to pull it into pieces. With this change, great durability is probably acquired; at least this is the case with the clasped petioles of Clematis vitalba. The meaning of these changes is obvious, namely, that the petioles may firmly and durably support the stem.
Clematis microphylla, var. leptophylla.--The long and thin internodes of this Australian species revolve sometimes in one direction and sometimes in an opposite one, describing long, narrow, irregular ellipses or large circles. Four revolutions were completed within five minutes of the same average rate of 1 hr. 51 m.; so that this species moves more quickly than the others of the genus. The shoots, when placed near a vertical stick, either twine round it, or clasp it with the basal portions of their petioles. The leaves whilst young are nearly of the same shape as those of C. viticella, and act in the same manner like a hook, as will be described under that species.
But the leaflets are more divided, and each segment whilst young terminates in a hardish point, which is much curved downwards and inwards; so that the whole leaf readily catches hold of any neighbouring object. The petioles of the young terminal leaflets are acted on by loops of thread weighing 0.125th and even 0.0625th of a grain. The basal portion of the main petiole is much less sensitive, but will clasp a stick against which it presses.
The leaves, whilst young, are continually and spontaneously moving slowly. A bell-gla.s.s was placed over a shoot secured to a stick, and the movements of the leaves were traced on it during several days. A very irregular line was generally formed; but one day, in the course of eight hours and three quarters, the figure clearly represented three and a half irregular ellipses, the most perfect one of which was completed in 2 hrs. 35 m. The two opposite leaves moved independently of each other. This movement of the leaves would aid that of the internodes in bringing the petioles into contact with surrounding objects. I discovered this movement too late to be enabled to observe it in the other species; but from a.n.a.logy I can hardly doubt that the leaves of at least C. viticella, C. flammula, and C. vitalba move spontaneously; and, judging from C Sieboldi, this probably is the case with C. montana and C. calycina. I ascertained that the simple leaves of C. glandulosa exhibited no spontaneous revolving movement.
Clematis viticella, var. venosa.--In this and the two following species the power of spirally twining is completely lost, and this seems due to the lessened flexibility of the internodes and to the interference caused by the large size of the leaves. But the revolving movement, though restricted, is not lost. In our present species a young internode, placed in front of a window, made three narrow ellipses, transversely to the direction of the light, at an average rate of 2 hrs. 40 m. When placed so that the movements were to and from the light, the rate was greatly accelerated in one half of the course, and r.e.t.a.r.ded in the other, as with twining plants.
The ellipses were small; the longer diameter, described by the apex of a shoot bearing a pair of not expanded leaves, was only 4.625 inches, and that by the apex of the penultimate internode only 1.125 inch. At the most favourable period of growth each leaf would hardly be carried to and fro by the movement of the internodes more than two or three inches, but, as above stated, it is probable that the leaves themselves move spontaneously. The movement of the whole shoot by the wind and by its rapid growth, would probably be almost equally efficient as these spontaneous movements, in bringing the petioles into contact with surrounding objects.
The leaves are of large size. Each bears three pairs of lateral leaflets and a terminal one, all supported on rather long sub- petioles. The main petiole bends a little angularly downwards at each point where a pair of leaflets arises (see fig. 2), and the petiole of the terminal leaflet is bent downwards at right angles; hence the whole petiole, with its rectangularly bent extremity, acts as a hook. This hook, the lateral petioles being directed a little upwards; forms an excellent grappling apparatus, by which the leaves readily become entangled with surrounding objects. If they catch nothing, the whole petiole ultimately grows straight. The main petiole, the sub-petioles, and the three branches into which each basi-lateral sub-petiole is generally subdivided, are all sensitive.
The basal portion of the main petiole, between the stem and the first pair of leaflets, is less sensitive than the remainder; it will, however, clasp a stick with which it is left in contact. The inferior surface of the rectangularly bent terminal portion (carrying the terminal leaflet), which forms the inner side of the end of the hook, is the most sensitive part; and this portion is manifestly best adapted to catch a distant support. To show the difference in sensibility, I gently placed loops of string of the same weight (in one instance weighing only 0.82 of a grain or 53.14 mg.) on the several lateral sub-petioles and on the terminal one; in a few hours the latter was bent, but after 24 hrs. no effect was produced on the other sub-petioles. Again, a terminal sub-petiole placed in contact with a thin stick became sensibly curved in 45 m., and in 1 hr. 10m.
moved through ninety degrees; whilst a lateral sub-petiole did not become sensibly curved until 3 hrs. 30 m. had elapsed. In all cases, if the sticks are taken away, the petioles continue to move during many hours afterwards; so they do after a slight rubbing; but they become straight again, after about a day"s interval, that is if the flexure has not been very great or long continued.
The graduated difference in the extension of the sensitiveness in the petioles of the above-described species deserves notice. In C.
montana it is confined to the main petiole, and has not spread to the sub-petioles of the three leaflets; so it is with young plants of C.
calycina, but in older plants it spreads to the three sub-petioles.
In C. viticella the sensitiveness has spread to the petioles of the seven leaflets, and to the subdivisions of the basi-lateral sub- petioles. But in this latter species it has diminished in the basal part of the main petiole, in which alone it resided in C. montana; whilst it has increased in the abruptly bent terminal portion.
Clematis flammula.--The rather thick, straight, and stiff shoots, whilst growing vigorously in the spring, make small oval revolutions, following the sun in their course. Four were made at an average rate of 3 hrs. 45 m. The longer axis of the oval, described by the extreme tip, was directed at right angles to the line joining the opposite leaves; its length was in one case only 1.375, and in another case 1.75 inch; so that the young leaves were moved a very short distance. The shoots of the same plant observed in midsummer, when growing not so quickly, did not revolve at all. I cut down another plant in the early summer, so that by August 1st it had formed new and moderately vigorous shoots; these, when observed under a bell-gla.s.s, were on some days quite stationary, and on other days moved to and fro only about the eighth of an inch. Consequently the revolving power is much enfeebled in this species, and under unfavourable circ.u.mstances is completely lost. The shoot must depend for coming into contact with surrounding objects on the probable, though not ascertained spontaneous movement of the leaves, on rapid growth, and on movement from the wind. Hence, perhaps, it is that the petioles have acquired a high degree of sensitiveness as a compensation for the little power of movement in the shoots.
The petioles are bowed downwards, and have the same general hook-like form as in C. viticella. The medial petiole and the lateral sub- petioles are sensitive, especially the much bent terminal portion.
As the sensitiveness is here greater than in any other species of the genus observed by me, and is in itself remarkable, I will give fuller details. The petioles, when so young that they have not separated from one another, are not sensitive; when the lamina of a leaflet has grown to a quarter of an inch in length (that is, about one-sixth of its full size), the sensitiveness is highest; but at this period the petioles are relatively much more fully developed than are the blades of the leaves. Full-grown petioles are not in the least sensitive.
A thin stick placed so as to press lightly against a petiole, having a leaflet a quarter of an inch in length, caused the petiole to bend in 3 hrs. 15 m. In another case a petiole curled completely round a stick in 12 hrs. These petioles were left curled for 24 hrs., and the sticks were then removed; but they never straightened themselves. I took a twig, thinner than the petiole itself, and with it lightly rubbed several petioles four times up and down; these in 1 hr. 45 m.
became slightly curled; the curvature increased during some hours and then began to decrease, but after 25 hrs. from the time of rubbing a vestige of the curvature remained. Some other petioles similarly rubbed twice, that is, once up and once down, became perceptibly curved in about 2 hrs. 30 m., the terminal sub-petiole moving more than the lateral sub-petioles; they all became straight again in between 12 hrs. and 14 hrs. Lastly, a length of about one-eighth of an inch of a sub-petiole, was lightly rubbed with the same twig only once; it became slightly curved in 3 hrs., remaining so during 11 hrs., but by the next morning was quite straight.
The following observations are more precise. After trying heavier pieces of string and thread, I placed a loop of fine string, weighing 1.04 gr. (67.4 mg.) on a terminal sub-petiole: in 6 hrs. 40 m. a curvature could be seen; in 24 hrs. the petiole formed an open ring round the string; in 48 hrs. the ring had almost closed on the string, and in 72 hrs. seized it so firmly, that some force was necessary for its withdrawal. A loop weighing 0.52 of a grain (33.7 mg.) caused in 14 hrs. a lateral sub-petiole just perceptibly to curve, and in 24 hrs. it moved through ninety degrees. These observations were made during the summer: the following were made in the spring, when the petioles apparently are more sensitive:- A loop of thread, weighing one-eighth of a grain (8.1 mg.), produced no effect on the lateral sub-petioles, but placed on a terminal one, caused it, after 24 hrs., to curve moderately; the curvature, though the loop remained suspended, was after 48 hrs. diminished, but never disappeared; showing that the petiole had become partially accustomed to the insufficient stimulus. This experiment was twice repeated with nearly the same result. Lastly, a loop of thread, weighing only one-sixteenth of a grain (4.05 mg.) was twice gently placed by a forceps on a terminal sub-petiole (the plant being, of course, in a still and closed room), and this weight certainly caused a flexure, which very slowly increased until the petiole moved through nearly ninety degrees: beyond this it did not move; nor did the petiole, the loop remaining suspended, ever become perfectly straight again.
When we consider, on the one hand, the thickness and stiffness of the petioles, and, on the other hand, the thinness and softness of fine cotton thread, and what an extremely small weight one-sixteenth of a grain (4.05 mg.) is, these facts are remarkable. But I have reason to believe that even a less weight excites curvature when pressing over a broader surface than that acted on by a thread. Having noticed that the end of a suspended string which accidentally touched a petiole, caused it to bend, I took two pieces of thin twine, 10 inches in length (weighing 1.64 gr.), and, tying them to a stick, let them hang as nearly perpendicularly downwards as their thinness and flexuous form, after being stretched, would permit; I then quietly placed their ends so as just to rest on two petioles, and these certainly became curved in 36 hrs. One of the ends touched the angle between a terminal and lateral sub-petiole, and it was in 48 hours caught between them as by a forceps. In these cases the pressure, though spread over a wider surface than that touched by the cotton thread, must have been excessively slight.
Clematis vitalba.--The plants were in pots and not healthy, so that I dare not trust my observations, which indicate much similarity in habits with C. flammula. I mention this species only because I have seen many proofs that the petioles in a state of nature are excited to movement by very slight pressure. For instance, I have found them embracing thin withered blades of gra.s.s, the soft young leaves of a maple, and the flower-peduncles of the quaking-gra.s.s or Briza. The latter are about as thick as the hair of a man"s beard, but they were completely surrounded and clasped. The petioles of a leaf, so young that none of the leaflets were expanded, had partially seized a twig.
Those of almost all the old leaves, even when unattached to any object, are much convoluted; but this is owing to their having come, whilst young, into contact during several hours with some object subsequently removed. With none of the above-described species, cultivated in pots and carefully observed, was there any permanent bending of the petioles without the stimulus of contact. In winter, the blades of the leaves of C. vitalba drop off; but the petioles (as was observed by Mohl) remain attached to the branches, sometimes during two seasons; and, being convoluted, they curiously resemble true tendrils, such as those possessed by the allied genus Naravelia.
The petioles which have clasped some object become much more stiff, hard, and polished than those which have failed in this their proper function.
TROPAEOLUM.--I observed T. tricolorum, T. azureum, T. pentaphyllum, T. peregrinum, T. elegans, T. tuberosum, and a dwarf variety of, as I believe, T. minus.
Tropaeolum tricolorum, var. grandiflorum.--The flexible shoots, which first rise from the tubers, are as thin as fine twine. One such shoot revolved in a course opposed to the sun, at an average rate, judging from three revolutions, of 1 hr. 23 m.; but no doubt the direction of the revolving movement is variable. When the plants have grown tall and are branched, all the many lateral shoots revolve. The stem, whilst young, twines regularly round a thin vertical stick, and in one case I counted eight spiral turns in the same direction; but when grown older, the stem often runs straight up for a s.p.a.ce, and, being arrested by the clasping petioles, makes one or two spires in a reversed direction. Until the plant grows to a height of two or three feet, requiring about a month from the time when the first shoot appears above ground, no true leaves are produced, but, in their place, filaments coloured like the stem. The extremities of these filaments are pointed, a little flattened, and furrowed on the upper surface. They never become developed into leaves. As the plant grows in height new filaments are produced with slightly enlarged tips; then others, bearing on each side of the enlarged medial tip a rudimentary segment of a leaf; soon other segments appear, and at last a perfect leaf is formed, with seven deep segments. So that on the same plant we may see every step, from tendril-like clasping filaments to perfect leaves with clasping petioles. After the plant has grown to a considerable height, and is secured to its support by the petioles of the true leaves, the clasping filaments on the lower part of the stem wither and drop off; so that they perform only a temporary service.
These filaments or rudimentary leaves, as well as the petioles of the perfect leaves, whilst young, are highly sensitive on all sides to a touch. The slightest rub caused them to curve towards the rubbed side in about three minutes, and one bent itself into a ring in six minutes; they subsequently became straight. When, however, they have once completely clasped a stick, if this is removed, they do not straighten themselves. The most remarkable fact, and one which I have observed in no other species of the genus, is that the filaments and the petioles of the young leaves, if they catch no object, after standing for some days in their original position, spontaneously and slowly oscillate a little from side to side, and then move towards the stem and clasp it. They likewise often become, after a time, in some degree spirally contracted. They therefore fully deserve to be called tendrils, as they are used for climbing, are sensitive to a touch, move spontaneously, and ultimately contract into a spire, though an imperfect one. The present species would have been cla.s.sed amongst the tendril-bearers, had not these characters been confined to early youth. During maturity it is a true leaf-climber.
Tropaeolum azureum.--An upper internode made four revolutions, following the sun, at an average rate of 1 hr. 47 m. The stem twined spirally round a support in the same irregular manner as that of the last species. Rudimentary leaves or filaments do not exist. The petioles of the young leaves are very sensitive: a single light rub with a twig caused one to move perceptibly in 5 m., and another in 6 m. The former became bent at right angles in 15 min., and became straight again in between 5 hrs. and 6 hrs. A loop of thread weighing 0.125th of a grain caused another petiole to curve.
Tropaeolum pentaphyllum.--This species has not the power of spirally twining, which seems due, not so much to a want of flexibility in the stem, as to continual interference from the clasping petioles. An upper internode made three revolutions, following the sun, at an average rate of 1 hr. 46 m. The main purpose of the revolving movement in all the species of Tropaeolum manifestly is to bring the petioles into contact with some supporting object. The petiole of a young leaf, after a slight rub, became curved in 6 m.; another, on a cold day, in 20 m., and others in from 8 m. to 10 m. Their curvature usually increased greatly in from 15 m. to 20 m., and they became straight again in between 5 hrs. and 6 hrs., but on one occasion in 3 hrs. When a petiole has fairly clasped a stick, it is not able, on the removal of the stick, to straighten itself. The free upper part of one, the base of which had already clasped a stick, still retained the power of movement. A loop of thread weighing 0.125th of a grain caused a petiole to curve; but the stimulus was not sufficient, the loop remaining suspended, to cause a permanent flexure. If a much heavier loop be placed in the angle between the petiole and the stem, it produces no effect; whereas we have seen with Clematis montana that the angle between the stem and petiole is sensitive.
Tropaeolum peregrinum.--The first-formed internodes of a young plant did not revolve, resembling in this respect those of a twining plant.