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1、【英文文学】The Movements and Habits of Climbing PlantsPrefaceThis Essay first appeared in the ninth volume of the Journal of the Linnean Society, published in 1865. It is here reproduced in a corrected and, I hope, clearer form, with some additional facts. The illustrations were drawn by my son, George D
2、arwin. Fritz Muller, after the publication of my paper, sent to the Linnean Society (Journal, vol. ix., p. 344) some interesting observations on the climbing plants of South Brazil, to which I shall frequently refer. Recently two important memoirs, chiefly on the difference in growth between the upp
3、er and lower sides of tendrils, and on the mechanism of the movements of twining-plants, by Dr. Hugo de Vries, have appeared in the Arbeiten des Botanischen Instituts in Wurzburg, Heft. iii., 1873. These memoirs ought to be carefully studied by every one interested in the subject, as I can here give
4、 only references to the more important points. This excellent observer, as well as Professor Sachs, 1 attributes all the movements of tendrils to rapid growth along one side; but, from reasons assigned towards the close of my fourth chapter, I cannot persuade myself that this holds good with respect
5、 to those due to a touch. In order that the reader may know what points have interested me most, I may call his attention to certain tendril-bearing plants; for instance, Bignonia capreolata, Cobaea, Echinocystis, and Hanburya, which display as beautiful adaptations as can be found in any part of th
6、e kingdom of nature. It is, also, an interesting fact that intermediate states between organs fitted for widely different functions, may be observed on the same individual plant of Corydalis claviculata and the common vine; and these cases illustrate in a striking manner the principle of the gradual
7、 evolution of species.Appendix to Preface (1882).Since the publication of this Edition two papers by eminent botanists have appeared; Schwendener, Das Winden der Pflanzen (Monatsberichte der Berliner Akademie, Dec. 1881), and J. Sachs, Notiz uber Schlingpflanzen (Arbeiten des botanischen Instituts i
8、n Wurzburg, Bd. ii. p. 719, 1882). The view “that the capacity of revolving, on which most climbers depend, is inherent, though undeveloped, in almost every plant in the vegetable kingdom” (Climbing Plants, p. 205), has been confirmed by the observations on circumnutation since given in The Power of
9、 Movement in Plants.Errata.On pp. 28, 32, 40, 53, statements are made with reference to the supposed acceleration of the revolving movement towards the light. It appears from the observations given in The Power of Movement in Plants, p. 451, that these conclusions were drawn from insufficient observ
10、ations, and are erroneous.Chapter I. Twining PlantsIntroductory remarks Description of the twining of the Hop Torsion of the stems Nature of the revolving movement, and manner of ascent Stems not irritable Rate of revolution in various plants Thickness of the support round which plants can twine Spe
11、cies which revolve in an anomalous manner.I was led to this subject by an interesting, but short paper by Professor Asa Gray on the movements of the tendrils of some Cucurbitaceous plants. 2 My observations were more than half completed before I learnt that the surprising phenomenon of the spontaneo
12、us revolutions of the stems and tendrils of climbing plants had been long ago observed by Palm and by Hugo von Mohl, 3 and had subsequently been the subject of two memoirs by Dutrochet. 4 Nevertheless, I believe that my observations, founded on the examination of above a hundred widely distinct livi
13、ng species, contain sufficient novelty to justify me in publishing them.Climbing plants may be divided into four classes. First, those which twine spirally round a support, and are not aided by any other movement. Secondly, those endowed with irritable organs, which when they touch any object clasp
14、it; such organs consisting of modified leaves, branches, or flower-peduncles. But these two classes sometimes graduate to a certain extent into one another. Plants of the third class ascend merely by the aid of hooks; and those of the fourth by rootlets; but as in neither class do the plants exhibit
15、 any special movements, they present little interest, and generally when I speak of climbing plants I refer to the two first great classes.TWINING PLANTS.This is the largest subdivision, and is apparently the primordial and simplest condition of the class. My observations will be best given by takin
16、g a few special cases. When the shoot of a Hop (Humulus lupulus) rises from the ground, the two or three first-formed joints or internodes are straight and remain stationary; but the next-formed, whilst very young, may be seen to bend to one side and to travel slowly round towards all points of the
17、compass, moving, like the hands of a watch, with the sun. The movement very soon acquires its full ordinary velocity. From seven observations made during August on shoots proceeding from a plant which had been cut down, and on another plant during April, the average rate during hot weather and durin
18、g the day is 2 hrs. 8 m. for each revolution; and none of the revolutions varied much from this rate. The revolving movement continues as long as the plant continues to grow; but each separate internode, as it becomes old, ceases to move.To ascertain more precisely what amount of movement each inter
19、node underwent, I kept a potted plant, during the night and day, in a well-warmed room to which I was confined by illness. A long shoot projected beyond the upper end of the supporting stick, and was steadily revolving. I then took a longer stick and tied up the shoot, so that only a very young inte
20、rnode, 1.75 of an inch in length, was left free. This was so nearly upright that its revolution could not be easily observed; but it certainly moved, and the side of the internode which was at one time convex became concave, which, as we shall hereafter see, is a sure sign of the revolving movement.
21、 I will assume that it made at least one revolution during the first twenty-four hours. Early the next morning its position was marked, and it made a second revolution in 9 hrs.; during the latter part of this revolution it moved much quicker, and the third circle was performed in the evening in a l
22、ittle over 3 hrs. As on the succeeding morning I found that the shoot revolved in 2 hrs. 45 m., it must have made during the night four revolutions, each at the average rate of a little over 3 hrs. I should add that the temperature of the room varied only a little. The shoot had now grown 3.5 inches
23、 in length, and carried at its extremity a young internode 1 inch in length, which showed slight changes in its curvature. The next or ninth revolution was effected in 2 hrs. 30 m. From this time forward, the revolutions were easily observed. The thirty-sixth revolution was performed at the usual ra
24、te; so was the last or thirty-seventh, but it was not completed; for the internode suddenly became upright, and after moving to the centre, remained motionless. I tied a weight to its upper end, so as to bow it slightly and thus detect any movement; but there was none. Some time before the last revo
25、lution was half performed, the lower part of the internode ceased to move.A few more remarks will complete all that need be said about this internode. It moved during five days; but the more rapid movements, after the performance of the third revolution, lasted during three days and twenty hours. Th
26、e regular revolutions, from the ninth to thirty-sixth inclusive, were effected at the average rate of 2 hrs. 31 m.; but the weather was cold, and this affected the temperature of the room, especially during the night, and consequently retarded the rate of movement a little. There was only one irregu
27、lar movement, which consisted in the stem rapidly making, after an unusually slow revolution, only the segment of a circle. After the seventeenth revolution the internode had grown from 1.75 to 6 inches in length, and carried an internode 1.875 inch long, which was just perceptibly moving; and this
28、carried a very minute ultimate internode. After the twenty-first revolution, the penultimate internode was 2.5 inches long, and probably revolved in a period of about three hours. At the twenty-seventh revolution the lower and still moving internode was 8.375, the penultimate 3.5, and the ultimate 2
29、.5 inches in length; and the inclination of the whole shoot was such, that a circle 19 inches in diameter was swept by it. When the movement ceased, the lower internode was 9 inches, and the penultimate 6 inches in length; so that, from the twenty-seventh to thirty-seventh revolutions inclusive, thr
30、ee internodes were at the same time revolving.The lower internode, when it ceased revolving, became upright and rigid; but as the whole shoot was left to grow unsupported, it became after a time bent into a nearly horizontal position, the uppermost and growing internodes still revolving at the extre
31、mity, but of course no longer round the old central point of the supporting stick. From the changed position of the centre of gravity of the extremity, as it revolved, a slight and slow swaying movement was given to the long horizontally projecting shoot; and this movement I at first thought was a s
32、pontaneous one. As the shoot grew, it hung down more and more, whilst the growing and revolving extremity turned itself up more and more.With the Hop we have seen that three internodes were at the same time revolving; and this was the case with most of the plants observed by me. With all, if in full
33、 health, two internodes revolved; so that by the time the lower one ceased to revolve, the one above was in full action, with a terminal internode just commencing to move. With Hoya carnosa, on the other hand, a depending shoot, without any developed leaves, 32 inches in length, and consisting of se
34、ven internodes (a minute terminal one, an inch in length, being counted), continually, but slowly, swayed from side to side in a semicircular course, with the extreme internodes making complete revolutions. This swaying movement was certainly due to the movement of the lower internodes, which, howev
35、er, had not force sufficient to swing the whole shoot round the central supporting stick. The case of another Asclepiadaceous plant, viz., Ceropegia Gardnerii, is worth briefly giving. I allowed the top to grow out almost horizontally to the length of 31 inches; this now consisted of three long inte
36、rnodes, terminated by two short ones. The whole revolved in a course opposed to the sun (the reverse of that of the Hop), at rates between 5 hrs. 15 m. and 6 hrs. 45 m. for each revolution. The extreme tip thus made a circle of above 5 feet (or 62 inches) in diameter and 16 feet in circumference, tr
37、avelling at the rate of 32 or 33 inches per hour. The weather being hot, the plant was allowed to stand on my study-table; and it was an interesting spectacle to watch the long shoot sweeping this grand circle, night and day, in search of some object round which to twine.If we take hold of a growing
38、 sapling, we can of course bend it to all sides in succession, so as to make the tip describe a circle, like that performed by the summit of a spontaneously revolving plant. By this movement the sapling is not in the least twisted round its own axis. I mention this because if a black point be painte
39、d on the bark, on the side which is uppermost when the sapling is bent towards the holders body, as the circle is described, the black point gradually turns round and sinks to the lower side, and comes up again when the circle is completed; and this gives the false appearance of twisting, which, in
40、the case of spontaneously revolving plants, deceived me for a time. The appearance is the more deceitful because the axes of nearly all twining-plants are really twisted; and they are twisted in the same direction with the spontaneous revolving movement. To give an instance, the internode of the Hop
41、 of which the history has been recorded, was at first, as could be seen by the ridges on its surface, not in the least twisted; but when, after the 37th revolution, it had grown 9 inches long, and its revolving movement had ceased, it had become twisted three times round its own axis, in the line of
42、 the course of the sun; on the other hand, the common Convolvulus, which revolves in an opposite course to the Hop, becomes twisted in an opposite direction.Hence it is not surprising that Hugo von Mohl (p. 105, 108, &c.) thought that the twisting of the axis caused the revolving movement; but it is
43、 not possible that the twisting of the axis of the Hop three times should have caused thirty-seven revolutions. Moreover, the revolving movement commenced in the young internode before any twisting of its axis could be detected. The internodes of a young Siphomeris and Lecontea revolved during sever
44、al days, but became twisted only once round their own axes. The best evidence, however, that the twisting does not cause the revolving movement is afforded by many leaf-climbing and tendril-bearing plants (as Pisum sativum, Echinocystis lobata, Bignonia capreolata, Eccremocarpus scaber, and with the
45、 leaf-climbers, Solanum jasminoides and various species of Clematis), of which the internodes are not twisted, but which, as we shall hereafter see, regularly perform revolving movements like those of true twining-plants. Moreover, according to Palm (pp. 30, 95) and Mohl (p. 149), and Leon, 5 intern
46、odes may occasionally, and even not very rarely, be found which are twisted in an opposite direction to the other internodes on the same plant, and to the course of their revolutions; and this, according to Leon (p. 356), is the case with all the internodes of a certain variety of Phaseolus multiflo
47、rus. Internodes which have become twisted round their own axes, if they have not ceased to revolve, are still capable of twining round a support, as I have several times observed.Mohl has remarked (p. 111) that when a stem twines round a smooth cylindrical stick, it does not become twisted. 6 Accord
48、ingly I allowed kidney-beans to run up stretched string, and up smooth rods of iron and glass, one-third of an inch in diameter, and they became twisted only in that degree which follows as a mechanical necessity from the spiral winding. The stems, on the other hand, which had ascended ordinary roug
49、h sticks were all more or less and generally much twisted. The influence of the roughness of the support in causing axial twisting was well seen in the stems which had twined up the glass rods; for these rods were fixed into split sticks below, and were secured above to cross sticks, and the stems in passing these places became much twisted. As soon as the stems which had ascended the iron rods reached the summit and became free, they also became twisted; and this apparently occurred more quick