The Movements and Habits of Climbing Plants · Charles Darwin
CHAPTER IV. TENDRIL-BEARERS—(_continued_).
In the course of about two days after a tendril has arranged its branches so as to press on any surface, the curved tips swell, become bright red, and form on their under-sides the well-known little discs or cushions with which they adhere firmly. In one case the tips were slightly swollen in 38 hrs. after coming into contact with a brick; in another case they were considerably swollen in 48 hrs., and in an additional 24 hrs. were firmly attached to a smooth board; and lastly, the tips of a younger tendril not only swelled but became attached to a stuccoed wall in 42 hrs. These adhesive discs resemble, except in colour and in being larger, those of _Bignonia capreolata_. When they were developed in contact with a ball of tow, the fibres were separately enveloped, but not in so effective a manner as by _B. capreolata_. Discs are never developed, as far as I have seen, without the stimulus of at least temporary contact with some object. {146} They are generally first formed on one side of the curved tip, the whole of which often becomes so much changed in appearance, that a line of the original green tissue can be traced only along the concave surface. When, however, a tendril has clasped a cylindrical stick, an irregular rim or disc is sometimes formed along the inner surface at some little distance from the curved tip; this was also observed (p. 71) by Mohl. The discs consist of enlarged cells, with smooth projecting hemispherical surfaces, coloured red; they are at first gorged with fluid (see section given by Mohl, p. 70), but ultimately become woody.
A tendril which has not become attached to any body, does not contract spirally; and in course of a week or two shrinks into the finest thread, withers and drops off. An attached tendril, on the other hand, contracts spirally, and thus becomes highly elastic, so that when the main foot-stalk is pulled the strain is distributed equally between all the attached discs. For a few days after the attachment of the discs, the tendril remains weak and brittle, but it rapidly increases in thickness and acquires great strength. During the following winter it ceases to live, but adheres firmly in a dead state both to its own stem and to the surface of attachment. In the accompanying diagram (fig. 11.) we see the difference between a tendril (B) some weeks after its attachment to a wall, with one (A) from the same plant fully grown but unattached. That the change in the nature of the tissues, as well as the spiral contraction, are consequent on the formation of the discs, is well shown by any lateral branches which have not become attached; for these in a week or two wither and drop off, in the same manner as does the whole tendril if unattached. The gain in strength and durability in a tendril after its attachment is something wonderful. There are tendrils now adhering to my house which are still strong, and have been exposed to the weather in a dead state for fourteen or fifteen years. One single lateral branchlet of a tendril, estimated to be at least ten years old, was still elastic and supported a weight of exactly two pounds. The whole tendril had five disc-bearing branches of equal thickness and apparently of equal strength; so that after having been exposed during ten years to the weather, it would probably have resisted a strain of ten pounds!
The two or three upper internodes, whilst young, steadily revolve; those on one plant made two circles, against the course of the sun, in 3 hrs. 12 m.; in a second plant the same course was followed, and the two circles were completed in 3 hrs. 41 m.; in a third plant, the internodes followed the sun and made two circles in 3 hrs. 47 m. The average rate of these six revolutions was 1 hr. 46 m. The stem shows no tendency to twine spirally round a support; but the allied tendril-bearing genus _Paullinia_ is said (Mohl, p. 4) to be a twiner. The flower-peduncles, which stand up above the end of the shoot, are carried round and round by the revolving movement of the internodes; and when the stem is securely tied, the long and thin flower-peduncles themselves are seen to be in continued and sometimes rapid movement from side to side. They sweep a wide space, but only occasionally revolve in a regular elliptical course. By the combined movements of the internodes and peduncles, one of the two short hooked tendrils, sooner or later, catches hold of some twig or branch, and then it curls round and securely grasps it.
These tendrils are, however, but slightly sensitive; for by rubbing their under surface only a slight movement is slowly produced. I hooked a tendril on to a twig; and in 1 hr. 45 m. it was curved considerably inwards; in 2 hrs. 30 m. it formed a ring; and in from 5 to 6 hours from being first hooked, it closely grasped the stick. A second tendril acted at nearly the same rate; but I observed one that took 24 hours before it curled twice round a thin twig. Tendrils which have caught nothing, spontaneously curl up to a close helix after the interval of several days. Those which have curled round some object, soon become a little thicker and tougher. The long and thin main peduncle, though spontaneously moving, is not sensitive and never clasps a support. Nor does it ever contract spirally, {152} although a contraction of this kind apparently would have been of service to the plant in climbing. Nevertheless it climbs pretty well without this aid. The seed-capsules though light, are of enormous size (hence its English name of balloon-vine), and as two or three are carried on the same peduncle, the tendrils rising close to them may be of service in preventing their being dashed to pieces by the wind. In the hothouse the tendrils served simply for climbing.
The position of the tendrils alone suffices to show their homological nature. In two instances one of two tendrils produced a flower at its tip; this, however, did not prevent its acting properly and curling round a twig. In a third case both lateral branches which ought to have been modified into tendrils, produced flowers like the central branch, and had quite lost their tendril-structure. I have seen, but was not enabled carefully to observe, only one other climbing Sapindaceous plant, namely, _Paullinia_. It was not in flower, yet bore long forked tendrils. So that, _Paullinia_, with respect to its tendrils, appears to bear the same relation to _Cardiospermum_ that _Cissus_ does to _Vitis_. PASSIFLORACEÆ.—After reading the discussion and facts given by Mohl (p. 47) on the nature of the tendrils in this family, no one can doubt that they are modified flower-peduncles. The tendrils and the flower-peduncles rise close side by side; and my son, William E. Darwin, made sketches for me of their earliest state of development in the hybrid _P. floribunda_. The two organs appear at first as a single papilla which gradually divides; so that the tendril appears to be a modified branch of the flower-peduncle. My son found one very young tendril surmounted by traces of floral organs, exactly like those on the summit of the true flower-peduncle at the same early age.
_Passiflora gracilis_.—This well-named, elegant, annual species differs from the other members of the group observed by me, in the young internodes having the power of revolving. It exceeds all the other climbing plants which I have examined, in the rapidity of its movements, and all tendril-bearers in the sensitiveness of the tendrils. The internode which carries the upper active tendril and which likewise carries one or two younger immature internodes, made three revolutions, following the sun, at an average rate of 1 hr. 4 m.; it then made, the day becoming very hot, three other revolutions at an average rate of between 57 and 58 m.; so that the average of all six revolutions was 1 hr. 1 m. The apex of the tendril describes elongated ellipses, sometimes narrow and sometimes broad, with their longer axes inclined in slightly different directions. The plant can ascend a thin upright stick by the aid of its tendrils; but the stem is too stiff for it to twine spirally round it, even when not interfered with by the tendrils, these having been successively pinched off at an early age.
When the stem is secured, the tendrils are seen to revolve in nearly the same manner and at the same rate as the internodes. {154} The tendrils are very thin, delicate, and straight, with the exception of the tips, which are a little curved; they are from 7 to 9 inches in length. A half-grown tendril is not sensitive; but when nearly full-grown they are extremely sensitive. A single delicate touch on the concave surface of the tip soon caused one to curve; and in 2 minutes it formed an open helix. A loop of soft thread weighing one thirty-second of a grain (2.02 mg.) placed most gently on the tip, thrice caused distinct curvature. A bent bit of thin platina wire weighing only fiftieth of a grain (1.23 mg.) twice produced the same effect; but this latter weight, when left suspended, did not suffice to cause a permanent curvature. These trials were made under a bell-glass, so that the loops of thread and wire were not agitated by the wind. The movement after a touch is very rapid: I took hold of the lower part of several tendrils, and then touched their concave tips with a thin twig and watched them carefully through a lens; the tips evidently began to bend after the following intervals—31, 25, 32, 31, 28, 39, 31, and 30 seconds; so that the movement was generally perceptible in half a minute after a touch; but on one occasion it was distinctly visible in 25 seconds. One of the tendrils which thus became bent in 31 seconds, had been touched two hours previously and had coiled into a helix; so that in this interval it had straightened itself and had perfectly recovered its irritability.
To ascertain how often the same tendril would become curved when touched, I kept a plant in my study, which from being cooler than the hot-house was not very favourable for the experiment. The extremity was gently rubbed four or five times with a thin stick, and this was done as often as it was observed to have become nearly straight again after having been in action; and in the course of 54 hrs. it answered to the stimulus 21 times, becoming each time hooked or spiral. On the last occasion, however, the movement was very slight, and soon afterwards permanent spiral contraction commenced. No trials were made during the night, so that the tendril would perhaps have answered a greater number of times to the stimulus; though, on the other hand, from having no rest it might have become exhausted from so many quickly repeated efforts.
I repeated the experiment made on the _Echinocystis_, and placed several plants of this _Passiflora_ so close together, that their tendrils were repeatedly dragged over each other; but no curvature ensued. I likewise repeatedly flirted small drops of water from a brush on many tendrils, and syringed others so violently that the whole tendril was dashed about, but they never became curved. The impact from the drops of water was felt far more distinctly on my hand than that from the loops of thread (weighing one thirty-second of a grain) when allowed to fall on it from a height, and these loops, which caused the tendrils to become curved, had been placed most gently on them. Hence it is clear, that the tendrils either have become habituated to the touch of other tendrils and drops of rain, or that they were from the first rendered sensitive only to prolonged though excessively slight pressure of solid objects, with the exclusion of that from other tendrils. To show the difference in the kind of sensitiveness in different plants and likewise to show the force of the syringe used, I may add that the lightest jet from it instantly caused the leaves of a _Mimosa_ to close; whereas the loop of thread weighing one thirty-second of a grain, when rolled into a ball and placed gently on the glands at the bases of the leaflets of the _Mimosa_, caused no action.
_Passiflora punctata_.—The internodes do not move, but the tendrils revolve regularly. A half-grown and very sensitive tendril made three revolutions, opposed to the course of the sun, in 3 hrs. 5 m., 2 hrs. 40 m. and 2 hrs. 50 m.; perhaps it might have travelled more quickly when nearly full-grown. A plant was placed in front of a window, and, as with twining stems, the light accelerated the movement of the tendril in one direction and retarded it in the other; the semicircle towards the light being performed in one instance in 15 m. less time and in a second instance in 20 m. less time than that required by the semicircle towards the dark end of the room. Considering the extreme tenuity of these tendrils, the action of the light on them is remarkable. The tendrils are long, and, as just stated, very thin, with the tip slightly curved or hooked. The concave side is extremely sensitive to a touch—even a single touch causing it to curl inwards; it subsequently straightened itself, and was again ready to act. A loop of soft thread weighing one fourteenth of a grain (4.625 mg.) caused the extreme tip to bend; another time I tried to hang the same little loop on an inclined tendril, but three times it slid off; yet this extraordinarily slight degree of friction sufficed to make the tip curl. The tendril, though so sensitive, does not move very quickly after a touch, no conspicuous movement being observable until 5 or 10 m. had elapsed. The convex side of the tip is not sensitive to a touch or to a suspended loop of thread. On one occasion I observed a tendril revolving with the convex side of the tip forwards, and in consequence it was not able to clasp a stick, against which it scraped; whereas tendrils revolving with the concave side forward, promptly seize any object in their path.
_Passiflora quadrangularis_.—This is a very distinct species. The tendrils are thick, long, and stiff; they are sensitive to a touch only on the concave surface towards the extremity. When a stick was placed so that the middle of the tendril came into contact with it, no curvature ensued. In the hothouse a tendril made two revolutions, each in 2 hrs. 22 m.; in a cool room one was completed in 3 hrs., and a second in 4 hrs. The internodes do not revolve; nor do those of the hybrid _P. floribunda_. _Tacsonia manicata_.—Here again the internodes do not revolve. The tendrils are moderately thin and long; one made a narrow ellipse in 5 hrs. 20 m., and the next day a broad ellipse in 5 hrs. 7 m. The extremity being lightly rubbed on the concave surface, became just perceptibly curved in 7 m., distinctly in 10 m., and hooked in 20 m. We have seen that the tendrils in the last three families, namely, the Vitaceæ, Sapindaceæ and Passifloraceæ, are modified flower-peduncles. This is likewise the case, according to De Candolle (as quoted by Mohl), with the tendrils of _Brunnichia_, one of the Polygonaceæ. In two or three species of _Modecca_, one of the Papayaceæ, the tendrils, as I hear from Prof. Oliver, occasionally bear flowers and fruit; so that they are axial in their nature.
_The Spiral Contraction of Tendrils_. This movement, which shortens the tendrils and renders them elastic, commences in half a day, or in a day or two after their extremities have caught some object. There is no such movement in any leaf-climber, with the exception of an occasional trace of it in the petioles of _Tropæolum tricolorum_. On the other hand, the tendrils of all tendril-bearing plants, contract spirally after they have caught an object with the following exceptions. Firstly, _Corydalis claviculata_, but then this plant might be called a leaf-climber. Secondly and thirdly, _Bignonia unguis_ with its close allies, and _Cardiospermum_; but their tendrils are so short that their contraction could hardly occur, and would be quite superfluous. Fourthly, _Smilaæ aspera_ offers a more marked exception, as its tendrils are moderately long. The tendrils of _Dicentra_, whilst the plant is young, are short and after attachment only become slightly flexuous; in older plants they are longer and then they contract spirally. I have seen no other exceptions to the rule that tendrils, after clasping with their extremities a support, undergo spiral contraction. When, however, the tendril of a plant of which the stem is immovably fixed, catches some fixed object, it does not contract, simply because it cannot; this, however, rarely occurs. In the common Pea the lateral branches alone contract, and not the central stem; and with most plants, such as the Vine, Passiflora, Bryony, the basal portion never forms a spire.
I have said that in _Corydalis claviculata_ the end of the leaf or tendril (for this part may be indifferently so called) does not contract into a spire. The branchlets, however, after they have wound round thin twigs, become deeply sinuous or zigzag. Moreover the whole end of the petiole or tendril, if it seizes nothing, bends after a time abruptly downwards and inwards, showing that its outer surface has gone on growing after the inner surface has ceased to grow. That growth is the chief cause of the spiral contraction of tendrils may be safely admitted, as shown by the recent researches of H. de Vries. I will, however, add one little fact in support of this conclusion. If the short, nearly straight portion of an attached tendril of _Passiflora gracilis_, (and, as I believe, of other tendrils,) between the opposed spires, be examined, it will be found to be transversely wrinkled in a conspicuous manner on the outside; and this would naturally follow if the outer side had grown more than the inner side, this part being at the same time forcibly prevented from becoming curved. So again the whole outer surface of a spirally wound tendril becomes wrinkled if it be pulled straight. Nevertheless, as the contraction travels from the extremity of a tendril, after it has been stimulated by contact with a support, down to the base, I cannot avoid doubting, from reasons presently to be given, whether the whole effect ought to be attributed to growth. An unattached tendril rolls itself up into a flat helix, as in the case of Cardiospermum, if the contraction commences at the extremity and is quite regular; but if the continued growth of the outer surface is a little lateral, or if the process begins near the base, the terminal portion cannot be rolled up within the basal portion, and the tendril then forms a more or less open spire. A similar result follows if the extremity has caught some object, and is thus held fast.
The tendrils of many kinds of plants, if they catch nothing, contract after an interval of several days or weeks into a spire; but in these cases the movement takes place after the tendril has lost its revolving power and hangs down; it has also then partly or wholly lost its sensibility; so that this movement can be of no use. The spiral contraction of unattached tendrils is a much slower process than that of attached ones. Young tendrils which have caught a support and are spirally contracted, may constantly be seen on the same stem with the much older unattached and uncontracted tendrils. In the _Echinocystis_ I have seen a tendril with the two lateral branches encircling twigs and contracted into beautiful spires, whilst the main branch which had caught nothing remained for many days straight. In this plant I once observed a main branch after it had caught a stick become spirally flexuous in 7 hrs., and spirally contracted in 18 hrs. Generally the tendrils of the _Echinocystis_ begin to contract in from 12 hrs. to 24 hrs. after catching some object; whilst unattached tendrils do not begin to contract until two or three or even more days after all revolving movement has ceased.
A full-grown tendril of _Passiflora quadrangularis_ which had caught a stick began in 8 hrs. to contract, and in 24 hrs. formed several spires; a younger tendril, only two-thirds grown, showed the first trace of contraction in two days after clasping a stick, and in two more days formed several spires. It appears, therefore, that the contraction does not begin until the tendril is grown to nearly its full length. Another young tendril of about the same age and length as the last did not catch any object; it acquired its full length in four days; in six additional days it first became flexuous, and in two more days formed one complete spire. This first spire was formed towards the basal end, and the contraction steadily but slowly progressed towards the apex; but the whole was not closely wound up into a spire until 21 days had elapsed from the first observation, that is, until 17 days after the tendril had grown to its full length.
The spiral contraction of tendrils is quite independent of their power of spontaneously revolving, for it occurs in tendrils, such as those of _Lathyrus grandiflorus_ and _Ampelopsis hederacea_, which do not revolve. It is not necessarily related to the curling of the tips round a support, as we see with the Ampelopsis and _Bignonia capreolata_, in which the development of adherent discs suffices to cause spiral contraction. Yet in some cases this contraction seems connected with the curling or clasping movement, due to contact with a support; for not only does it soon follow this act, but the contraction generally begins close to the curled extremity, and travels downwards to the base. If, however, a tendril be very slack, the whole length almost simultaneously becomes at first flexuous and then spiral. Again, the tendrils of some few plants never contract spirally unless they have first seized hold of some object; if they catch nothing they hang down, remaining straight, until they wither and drop off: this is the case with the tendrils of Bignonia, which consist of modified leaves, and with those of three genera of the Vitaceæ, which are modified flower-peduncles. But in the great majority of cases, tendrils which have never come in contact with any object, after a time contract spirally. All these facts taken together, show that the act of clasping a support and the spiral contraction of the whole length of the tendril, are phenomena not necessarily connected.
The spiral contraction which ensues after a tendril has caught a support is of high service to the plant; hence its almost universal occurrence with species belonging to widely different orders. When a shoot is inclined and its tendril has caught an object above, the spiral contraction drags up the shoot. When the shoot is upright, the growth of the stem, after the tendrils have seized some object above, would leave it slack, were it not for the spiral contraction which draws up the stem as it increases in length. Thus there is no waste of growth, and the stretched stem ascends by the shortest course. When a terminal branchlet of the tendril of Cobæa catches a stick, we have seen how well the spiral contraction successively brings the other branchlets, one after the other, into contact with the stick, until the whole tendril grasps it in an inextricable knot. When a tendril has caught a yielding object, this is sometimes enveloped and still further secured by the spiral folds, as I have seen with _Passiflora quadrangularis_; but this action is of little importance.
Text from Project Gutenberg, public domain in the United States.