POWER OF MOVEMENT
POWER OF MOVEMENT
BY CHARLES DARWIN, LL.D., F.R.S.
THE CIRCUMNUTATING MOVEMENTS OF SEEDLING PLANTS.
Brassica oleracea, circumnutation of the radicle, of the arched hypocotyl whilst still buried beneath the ground, whilst rising above the ground and straightening itself, and when erect—Circumnutation of the cotyledons— Rate of movement—Analogous observations on various organs in species of Githago, Gossypium, Oxalis, Tropaeolum, Citrus, Aesculus, of several Leguminous and Cucurbitaceous genera, Opuntia, Helianthus, Primula, Cyclamen, Stapelia, Cerinthe, Nolana, Solanum, Beta, Ricinus, Quercus, Corylus, Pinus, Cycas, Canna, Allium, Asparagus, Phalaris, Zea, Avena, Nephrodium, and Selaginella...10-66
GENERAL CONSIDERATIONS ON THE MOVEMENTS AND GROWTH OF SEEDLING PLANTS.
Generality of the circumnutating movement—Radicles, their circumnutation of service—Manner in which they penetrate the ground—Manner in which hypocotyls and other organs break through the ground by being arched— Singular manner of germination in Megarrhiza, etc.—Abortion of cotyledons- -Circumnutation of hypocotyls and epicotyls whilst still buried and arched- -Their power of straightening themselves—Bursting of the seed-coats— Inherited effect of the arching process in hypo- [page vi.] gean hypocotyls—Circumnutation of hypocotyls and epicotyls when erect— Circumnutation of cotyledons—Pulvini or joints of cotyledons, duration of their activity, rudimentary in Oxalis corniculata, their development— Sensitiveness of cotyledons to light and consequent disturbance of their periodic movements—Sensitiveness of cotyledons to contact...Page 67-128
SENSITIVENESS OF THE APEX OF THE RADICLE TO CONTACT AND TO OTHER IRRITANTS.
Manner in which radicles bend when they encounter an obstacle in the soil— Vicia faba, tips of radicles highly sensitive to contact and other irritants—Effects of too high a temperature—Power of discriminating between objects attached on opposite sides—Tips of secondary radicles sensitive—Pisum, tips of radicles sensitive—Effects of such sensitiveness in overcoming geotropism—Secondary radicles—Phaseolus, tips of radicles hardly sensitive to contact, but highly sensitive to caustic and to the removal of a slice—Tropaeolum—Gossypium—Cucurbita—Raphanus—Aesculus, tip not sensitive to slight contact, highly sensitive to caustic—Quercus, tip highly sensitive to contact—Power of discrimination—Zea, tip highly sensitive, secondary radicles—Sensitiveness of radicles to moist air— Summary of chapter...129-200
THE CIRCUMNUTATING MOVEMENTS OF THE SEVERAL PARTS OF MATURE PLANTS.
Circumnutation of stems: concluding remarks on—Circumnutation of stolons: aid thus afforded in winding amongst the stems of surrounding plants— Circumnutation of flower-stems—Circumnutation of Dicotyledonous leaves— Singular oscillatory movement of leaves of Dionaea—Leaves of Cannabis sink at night—Leaves of Gymnosperms—Of Monocotyledons—Cryptogams—Concluding remarks on the circumnutation of leaves; generally rise in the evening and sink in the morning...201-262 [page vii.]
MODIFIED CIRCUMNUTATION: CLIMBING PLANTS; EPINASTIC AND HYPONASTIC MOVEMENTS.
Circumnutation modified through innate causes or through the action of external conditions—Innate causes—Climbing plants; similarity of their movements with those of ordinary plants; increased amplitude; occasional points of difference—Epinastic growth of young leaves—Hyponastic growth of the hypocotyls and epicotyls of seedlings—Hooked tips of climbing and other plants due to modified circumnutation—Ampelopsis tricuspidata— Smithia Pfundii—Straightening of the tip due to hyponasty—Epinastic growth and circumnutation of the flower-peduncles of Trifolium repens and Oxalis carnosa...Page 263-279
MODIFIED CIRCUMNUTATION: SLEEP OR NYCTITROPIC MOVEMENTS, THEIR USE: SLEEP OF COTYLEDONS.
Preliminary sketch of the sleep or nyctitropic movements of leaves— Presence of pulvini—The lessening of radiation the final cause of nyctitropic movements—Manner of trying experiments on leaves of Oxalis, Arachis, Cassia, Melilotus, Lotus and Marsilea and on the cotyledons of Mimosa—Concluding remarks on radiation from leaves—Small differences in the conditions make a great difference in the result - Description of the nyctitropic position and movements of the cotyledons of various plants— List of species—Concluding remarks—Independence of the nyctitropic movements of the leaves and cotyledons of the same species—Reasons for believing that the movements have been acquired for a special purpose...280-316
MODIFIED CIRCUMNUTATION: NYCTITROPIC OR SLEEP MOVEMENTS OF LEAVES.
Conditions necessary for these movements—List of Genera and Families, which include sleeping plants—Description of the movements in the several Genera—Oxalis: leaflets folded at [page viii.] night—Averrhoa: rapid movements of the leaflets—Porlieria: leaflets close when plant kept very dry—Tropaeolum: leaves do not sleep unless well illuminated during day—Lupinus: various modes of sleeping—Melilotus: singular movements of terminal leaflet—Trifolium—Desmodium: rudimentary lateral leaflets, movements of, not developed on young plants, state of their pulvini—Cassia: complex movements of the leaflets—Bauhinia: leaves folded at night—Mimosa pudica: compounded movements of leaves, effect of darkness—Mimosa albida, reduced leaflets of—Schrankia: downward movement of the pinnae—Marsilea: the only cryptogam known to sleep—Concluding remarks and summary—Nyctitropism consists of modified circumnutation, regulated by the alternations of light and darkness—Shape of first true leaves...Page 317-417
MODIFIED CIRCUMNUTATION: MOVEMENTS EXCITED BY LIGHT.
Distinction between heliotropism and the effects of light on the periodicity of the movements of leaves—Heliotropic movements of Beta, Solanum, Zea, and Avena—Heliotropic movements towards an obscure light in Apios, Brassica, Phalaris, Tropaeolum, and Cassia—Apheliotropic movements of tendrils of Bignonia—Of flower-peduncles of Cyclamen—Burying of the pods—Heliotropism and apheliotropism modified forms of circumnutation— Steps by which one movement is converted into the other— Transversal-heliotropismus or diaheliotropism influenced by epinasty, the weight of the part and apogeotropism—Apogeotropism overcome during the middle of the day by diaheliotropism—Effects of the weight of the blades of cotyledons—So called diurnal sleep—Chlorophyll injured by intense light—Movements to avoid intense light...418-448
SENSITIVENESS OF PLANTS TO LIGHT: ITS TRANSMITTED EFFECTS.
Uses of heliotropism—Insectivorous and climbing plants not heliotropic— Same organ heliotropic at one age and not at another—Extraordinary sensitiveness of some plants to light—The effects [page ix.] of light do not correspond with its intensity—Effects of previous illumination—Time required for the action of light—After-effects of light—Apogeotropism acts as soon as light fails—Accuracy with which plants bend to the light—This dependent on the illumination of one whole side of the part—Localised sensitiveness to light and its transmitted effects—Cotyledons of Phalaris, manner of bending—Results of the exclusion of light from their tips—Effects transmitted beneath the surface of the ground—Lateral illumination of the tip determines the direction of the curvature of the base—Cotyledons of Avena, curvature of basal part due to the illumination of upper part—Similar results with the hypocotyls of Brassica and Beta—Radicles of Sinapis apheliotropic, due to the sensitiveness of their tips—Concluding remarks and summary of chapter— Means by which circumnutation has been converted into heliotropism or apheliotropism...Page 449-492
MODIFIED CIRCUMNUTATION: MOVEMENTS EXCITED BY GRAVITATION.
Means of observation—Apogeotropism—Cytisus—Verbena—Beta—Gradual conversion of the movement of circumnutation into apogeotropism in Rubus, Lilium, Phalaris, Avena, and Brassica—Apogeotropism retarded by heliotropism—Effected by the aid of joints or pulvini—Movements of flower-peduncles of Oxalis—General remarks on apogeotropism—Geotropism— Movements of radicles—Burying of seed-capsules—Use of process—Trifolium subterraneum—Arachis—Amphicarpaea—Diageotropism—Conclusion...493-522
LOCALISED SENSITIVENESS TO GRAVITATION, AND ITS TRANSMITTED EFFECTS.
General considerations—Vicia faba, effects of amputating the tips of the radicles—Regeneration of the tips—Effects of a short exposure of the tips to geotropic action and their subsequent amputation—Effects of amputating the tips obliquely—Effects of cauterising the tips—Effects of grease on the tips—Pisum [page x.] sativum, tips of radicles cauterised transversely, and on their upper and lower sides—Phaseolus, cauterisation and grease on the tips—Gossypium— Cucurbita, tips cauterised transversely, and on their upper and lower sides—Zea, tips cauterised—Concluding remarks and summary of chapter— Advantages of the sensibility to geotropism being localised in the tips of the radicles...Page 523-545
SUMMARY AND CONCLUDING REMARKS.
Nature of the circumnutating movement—History of a germinating seed—The radicle first protrudes and circumnutates—Its tip highly sensitive— Emergence of the hypocotyl or of the epicotyl from the ground under the form of an arch—Its circumnutation and that of the cotyledons—The seedling throws up a leaf-bearing stem—The circumnutation of all the parts or organs—Modified circumnutation—Epinasty and hyponasty—Movements of climbing plants—Nyctitropic movements—Movements excited by light and gravitation—Localised sensitiveness—Resemblance between the movements of plants and animals—The tip of the radicle acts like a brain...546-573
THE MOVEMENTS OF PLANTS.
THE chief object of the present work is to describe and connect together several large classes of movement, common to almost all plants. The most widely prevalent movement is essentially of the same nature as that of the stem of a climbing plant, which bends successively to all points of the compass, so that the tip revolves. This movement has been called by Sachs "revolving nutation;" but we have found it much more convenient to use the terms circumnutation and circumnutate. As we shall have to say much about this movement, it will be useful here briefly to describe its nature. If we observe a circumnutating stem, which happens at the time to be bent, we will say towards the north, it will be found gradually to bend more and more easterly, until it faces the east; and so onwards to the south, then to the west, and back again to the north. If the movement had been quite regular, the apex would have described a circle, or rather, as the stem is always growing upwards, a circular spiral. But it generally describes irregular elliptical or oval figures; for the apex, after pointing in any one direction, commonly moves back to the opposite side, not, however, returning along the same line. Afterwards other irregular ellipses or ovals are successively described, with their longer [page 2] axes directed to different points of the compass. Whilst describing such figures, the apex often travels in a zigzag line, or makes small subordinate loops or triangles. In the case of leaves the ellipses are generally narrow.
Until recently the cause of all such bending movements was believed to be due to the increased growth of the side which becomes for a time convex; that this side does temporarily grow more quickly than the concave side has been well established; but De Vries has lately shown that such increased growth follows a previously increased state of turgescence on the convex side.* In the case of parts provided with a so-called joint, cushion or pulvinus, which consists of an aggregate of small cells that have ceased to increase in size from a very early age, we meet with similar movements; and here, as Pfeffer has shown** and as we shall see in the course of this work, the increased turgescence of the cells on opposite sides is not followed by increased growth. Wiesner denies in certain cases the accuracy of De Vries' conclusion about turgescence, and maintains*** that the increased extensibility of the cell-walls is the more important element. That such extensibility must accompany increased turgescence in order that the part may bend is manifest, and this has been insisted on by several botanists; but in the case of unicellular plants it can hardly fail to be the more important element. On the whole we may at present conclude that in-
* Sachs first showed ('Lehrbuch,' etc., 4th edit. p. 452) the intimate connection between turgescence and growth. For De Vries' interesting essay, 'Wachsthumskrmmungen mehrzelliger Organe,' see 'Bot. Zeitung,' Dec. 19, 1879, p. 830.
** 'Die Periodischen Bewegungen der Blattorgane,' 1875.
*** 'Untersuchungen ber den Heliotropismus,' Sitzb. der K. Akad. der Wissenschaft. (Vienna), Jan. 1880.
[page 3] creased growth, first on one side and then on another, is a secondary effect, and that the increased turgescence of the cells, together with the extensibility of their walls, is the primary cause of the movement of circumnutation.*
In the course of the present volume it will be shown that apparently every growing part of every plant is continually circumnutating, though often on a small scale. Even the stems of seedlings before they have broken through the ground, as well as their buried radicles, circumnutate, as far as the pressure of the surrounding earth permits. In this universally present movement we have the basis or groundwork for the acquirement, according to the requirements of the plant, of the most diversified movements. Thus, the great sweeps made by the stems of twining plants, and by the tendrils of other climbers, result from a mere increase in the amplitude of the ordinary movement of circumnutation. The position which young leaves and other organs ultimately assume is acquired by the circumnutating movement being increased in some one direction. the leaves of various plants are said to sleep at night, and it will be seen that their blades then assume a vertical position through modified circumnutation, in order to protect their upper surfaces from being chilled through radiation. The movements of various organs to the light, which are so general throughout the vegetable kingdom, and occasionally from the light, or transversely with respect to it, are all modified
* See Mr. Vines' excellent discussion ('Arbeiten des Bot. Instituts in Wrzburg,' B. II. pp. 142, 143, 1878) on this intricate subject. Hofmeister's observations ('Jahreschrifte des Vereins fr Vaterl. Naturkunde in Wrtemberg,' 1874, p. 211) on the curious movements of Spirogyra, a plant consisting of a single row of cells, are valuable in relation to this subject.
[page 4] forms of circumnutation; as again are the equally prevalent movements of stems, etc., towards the zenith, and of roots towards the centre of the earth. In accordance with these conclusions, a considerable difficulty in the way of evolution is in part removed, for it might have been asked, how did all these diversified movements for the most different purposes first arise? As the case stands, we know that there is always movement in progress, and its amplitude, or direction, or both, have only to be modified for the good of the plant in relation with internal or external stimuli.
Besides describing the several modified forms of circumnutation, some other subjects will be discussed. The two which have interested us most are, firstly, the fact that with some seedling plants the uppermost part alone is sensitive to light, and transmits an influence to the lower part, causing it to bend. If therefore the upper part be wholly protected from light, the lower part may be exposed for hours to it, and yet does not become in the least bent, although this would have occurred quickly if the upper part had been excited by light. Secondly, with the radicles of seedlings, the tip is sensitive to various stimuli, especially to very slight pressure, and when thus excited, transmits an influence to the upper part, causing it to bend from the pressed side. On the other hand, if the tip is subjected to the vapour of water proceeding from one side, the upper part of the radicle bends towards this side. Again it is the tip, as stated by Ciesielski, though denied by others, which is sensitive to the attraction of gravity, and by transmission causes the adjoining parts of the radicle to bend towards the centre of the earth. These several cases of the effects of contact, other irritants, vapour, light, and the [page 5] attraction of gravity being transmitted from the excited part for some little distance along the organ in question, have an important bearing on the theory of all such movements.
[Terminology.—A brief explanation of some terms which will be used, must here be given. With seedlings, the stem which supports the cotyledons (i.e. the organs which represent the first leaves) has been called by many botanists the hypocotyledonous stem, but for brevity sake we will speak of it merely as the hypocotyl: the stem immediately above the cotyledons will be called the epicotyl or plumule. The radicle can be distinguished from the hypocotyl only by the presence of root-hairs and the nature of its covering. The meaning of the word circumnutation has already been explained. Authors speak of positive and negative heliotropism,*—that is, the bending of an organ to or from the light; but it is much more convenient to confine the word heliotropism to bending towards the light, and to designate as apheliotropism bending from the light. There is another reason for this change, for writers, as we have observed, occasionally drop the adjectives positive and negative, and thus introduce confusion into their discussions. Diaheliotropism may express a position more or less transverse to the light and induced by it. In like manner positive geotropism, or bending towards the centre of the earth, will be called by us geotropism; apogeotropism will mean bending in opposition to gravity or from the centre of the earth; and diageotropism, a position more or less transverse to the radius of the earth. The words heliotropism and geotropism properly mean the act of moving in relation to the light or the earth; but in the same manner as gravitation, though defined as "the act of tending to the centre," is often used to express the cause of a body falling, so it will be found convenient occasionally to employ heliotropism and geotropism, etc., as the cause of the movements in question.
The term epinasty is now often used in Germany, and implies that the upper surface of an organ grows more quickly than the
* The highly useful terms of Heliotropism and Geotropism were first used by Dr. A. B. Frank: see his remarkable 'Beitrge zur Pflanzenphysiologie,' 1868. [page 6] lower surface, and thus causes it to bend downwards. Hyponasty is the reverse, and implies increased growth along the lower surface, causing the part to bend upwards.*
Methods of Observation.—The movements, sometimes very small and sometimes considerable in extent, of the various organs observed by us, were traced in the manner which after many trials we found to be best, and which must be described. Plants growing in pots were protected wholly from the light, or had light admitted from above, or on one side as the case might require, and were covered above by a large horizontal sheet of glass, and with another vertical sheet on one side. A glass filament, not thicker than a horsehair, and from a quarter to three-quarters of an inch in length, was affixed to the part to be observed by means of shellac dissolved in alcohol. The solution was allowed to evaporate, until it became so thick that it set hard in two or three seconds, and it never injured the tissues, even the tips of tender radicles, to which it was applied. To the end of the glass filament an excessively minute bead of black sealing-wax was cemented, below or behind which a bit of card with a black dot was fixed to a stick driven into the ground. The weight of the filament was so slight that even small leaves were not perceptibly pressed down. another method of observation, when much magnification of the movement was not required, will presently be described. The bead and the dot on the card were viewed through the horizontal or vertical glass-plate (according to the position of the object), and when one exactly covered the other, a dot was made on the glass-plate with a sharply pointed stick dipped in thick Indian-ink. Other dots were made at short intervals of time and these were afterwards joined by straight lines. The figures thus traced were therefore angular; but if dots had been made every 1 or 2 minutes, the lines would have been more curvilinear, as occurred when radicles were allowed to trace their own courses on smoked glass-plates. To make the dots accurately was the sole difficulty, and required some practice. Nor could this be done quite accurately, when the movement was much magnified, such as 30 times and upwards; yet even in this case the general course may be trusted. To test the accuracy of the above method of observation, a filament was fixed to an
* These terms are used in the sense given them by De Vries, 'Wrzburg Arbeiten,' Heft ii 1872, p. 252.
[page 7] inanimate object which was made to slide along a straight edge and dots were repeatedly made on a glass-plate; when these were joined, the result ought to have been a perfectly straight line, and the line was very nearly straight. It may be added that when the dot on the card was placed half-an-inch below or behind the bead of sealing-wax, and when the glass-plate (supposing it to have been properly curved) stood at a distance of 7 inches in front (a common distance), then the tracing represented the movement of the bead magnified 15 times.
Whenever a great increase of the movement was not required, another, and in some respects better, method of observation was followed. This consisted in fixing two minute triangles of thin paper, about 1/20 inch in height, to the two ends of the attached glass filament; and when their tips were brought into a line so that they covered one another, dots were made as before on the glass-plate. If we suppose the glass-plate to stand at a distance of seven inches from the end of the shoot bearing the filament, the dots when joined, will give nearly the same figure as if a filament seven inches long, dipped in ink, had been fixed to the moving shoot, and had inscribed its own course on the plate. The movement is thus considerably magnified; for instance, if a shoot one inch in length were bending, and the glass-plate stood at the distance of seven inches, the movement would be magnified eight times. It would, however, have been very difficult to have ascertained in each case how great a length of the shoot was bending; and this is indispensable for ascertaining the degree to which the movement is magnified.
After dots had been made on the glass-plates by either of the above methods, they were copied on tracing paper and joined by ruled lines, with arrows showing the direction of the movement. The nocturnal courses are represented by straight broken lines. the first dot is always made larger than the others, so as to catch the eye, as may be seen in the diagrams. The figures on the glass-plates were often drawn on too large a scale to be reproduced on the pages of this volume, and the proportion in which they have been reduced is always given.* Whenever it could be approximately told how much the movement had been magnified, this is stated. We have perhaps
* We are much indebted to Mr. Cooper for the care with which he has reduced and engraved our diagrams.
[page 8] introduced a superfluous number of diagrams; but they take up less space than a full description of the movements. Almost all the sketches of plants asleep, etc., were carefully drawn for us by Mr. George Darwin.
As shoots, leaves, etc., in circumnutating bend more and more, first in one direction and then in another, they were necessarily viewed at different times more or less obliquely; and as the dots were made on a flat surface, the apparent amount of movement is exaggerated according to the degree of obliquity of the point of view. It would, therefore, have been a much better plan to have used hemispherical glasses, if we had possessed them of all sizes, and if the bending part of the shoot had been distinctly hinged and could have been placed so as to have formed one of the radii of the sphere. But even in this case it would have been necessary afterwards to have projected the figures on paper; so that complete accuracy could not have been attained. From the distortion of our figures, owing to the above causes, they are of no use to any one who wishes to know the exact amount of movement, or the exact course pursued; but they serve excellently for ascertaining whether or not the part moved at all, as well as the general character of the movement.]
In the following chapters, the movements of a considerable number of plants are described; and the species have been arranged according to the system adopted by Hooker in Le Maout and Decaisne's 'Descriptive Botany.' No one who is not investigating the present subject need read all the details, which, however, we have thought it advisable to give. To save the reader trouble, the conclusions and most of the more important parts have been printed in larger type than the other parts. He may, if he thinks fit, read the last chapter first, as it includes a summary of the whole volume; and he will thus see what points interest him, and on which he requires the full evidence.
Finally, we must have the pleasure of returning our [page 9] sincere thanks to Sir Joseph Hooker and to Mr. W. Thiselton Dyer for their great kindness, in not only sending us plants from Kew, but in procuring others from several sources when they were required for our observations; also, for naming many species, and giving us information on various points. [page 10]
THE CIRCUMNUTATING MOVEMENTS OF SEEDLING PLANTS.
Brassica oleracea, circumnutation of the radicle, of the arched hypocotyl whilst still buried beneath the ground, whilst rising above the ground and straightening itself, and when erect—Circumnutation of the cotyledons— Rate of movement—Analogous observations on various organs in species of Githago, Gossypium, Oxalis, Tropaeolum, Citrus, Aesculus, of several Leguminous and Cucurbitaceous genera, Opuntia, Helianthus, Primula, Cyclamen, Stapelia, Cerinthe, Nolana, Solanum, Beta, Ricinus, Quercus, Corylus, Pinus, Cycas, Canna, Allium, Asparagus, Phalaris, Zea, Avena, Nephrodium, and Selaginella.
THE following chapter is devoted to the circumnutating movements of the radicles, hypocotyls, and cotyledons of seedling plants; and, when the cotyledons do not rise above the ground, to the movements of the epicotyl. But in a future chapter we shall have to recur to the movements of certain cotyledons which sleep at night.
[Brassica oleracea (Cruciferae)'.—Fuller details will be given with respect to the movements in this case than in any other, as space and time will thus ultimately be saved.
Radicle.—A seed with the radicle projecting .05 inch was fastened with shellac to a little plate of zinc, so that the radicle stood up vertically; and a fine glass filament was then fixed near its base, that is, close to the seed-coats. The seed was surrounded by little bits of wet sponge, and the movement of the bead at the end of the filament was traced (Fig. 1) during sixty hours. In this time the radicle increased in length from .05 to .11 inch. Had the filament been attached at first close to the apex of the radicle, and if it could have remained there all the time, the movement exhibited would have [page 11] been much greater, for at the close of our observations the tip, instead of standing vertically upwards, had become bowed downwards through geotropism, so as almost to touch the zinc plate. As far as we could roughly ascertain by measurements made with compasses on other seeds, the tip alone, for a length of only 2/100 to 3/100 of an inch, is acted on by geotropism. But the tracing shows that the basal part of the radicle continued to circumnutate irregularly during the whole time. The actual extreme amount of movement of the bead at the end of the filament was nearly .05 inch, but to what extent the movement of the radicle was magnified by the filament, which was nearly 3/4 inch in length, it was impossible to estimate.
Fig. 1. Brassica oleracea: circumnutation of radicle, traced on horizontal glass, from 9 A.M. Jan. 31st to 9 P.M. Feb. 2nd. Movement of bead at end of filament magnified about 40 times.
Another seed was treated and observed in the same manner, but the radicle in this case protruded .1 inch, and was not Fig. 2. Brassica oleracea: circumnutating and geotropic movement of radicle, traced on horizontal glass during 46 hours.
fastened so as to project quite vertically upwards. The filament was affixed close to its base. The tracing (Fig. 2, reduced by half) shows the movement from 9 A.M. Jan. 31st to 7 A.M. Feb. 2nd; but it continued to move during the whole of the [page 12] 2nd in the same general direction, and in a similar zigzag manner. From the radicle not being quite perpendicular when the filament was affixed geotropism came into play at once; but the irregular zigzag course shows that there was growth (probably preceded by turgescence), sometimes on one and sometimes on another side. Occasionally the bead remained stationary for about an hour, and then probably growth occurred on the side opposite to that which caused the geotropic curvature. In the case previously described the basal part of the very short radicle from being turned vertically upwards, was at first very little affected by geotropism. Filaments were affixed in two other instances to rather longer radicles protruding obliquely from seeds which had been turned upside down; and in these cases the lines traced on the horizontal glasses were only slightly zigzag, and the movement was always in the same general direction, through the action of geotropism. All these observations are liable to several causes of error, but we believe, from what will hereafter be shown with respect to the movements of the radicles of other plants, that they may be largely trusted.
Hypocotyl.—The hypocotyl protrudes through the seed-coats as a rectangular projection, which grows rapidly into an arch like the letter U turned upside down; the cotyledons being still enclosed within the seed. In whatever position the seed may be embedded in the earth or otherwise fixed, both legs of the arch bend upwards through apogeotropism, and thus rise vertically above the ground. As soon as this has taken place, or even earlier, the inner or concave surface of the arch grows more quickly than the upper or convex surface; and this tends to separate the two legs and aids in drawing the cotyledons out of the buried seed-coats. By the growth of the whole arch the cotyledons are ultimately dragged from beneath the ground, even from a considerable depth; and now the hypocotyl quickly straightens itself by the increased growth of the concave side.
Even whilst the arched or doubled hypocotyl is still beneath the ground, it circumnutates as much as the pressure of the surrounding soil will permit; but this was difficult to observe, because as soon as the arch is freed from lateral pressure the two legs begin to separate, even at a very early age, before the arch would naturally have reached the surface. Seeds were allowed to germinate on the surface of damp earth, and after they had fixed themselves by their radicles, and after the, as yet, only [page 13] slightly arched hypocotyl had become nearly vertical, a glass filament was affixed on two occasions near to the base of the basal leg (i.e. the one in connection with the radicle), and its movements were traced in darkness on a horizontal glass. The result was that long lines were formed running in nearly the plane of the vertical arch, due to the early separation of the two legs now freed from pressure; but as the lines were zigzag, showing lateral movement, the arch must have been circumnutating, whilst it was straightening itself by growth along its inner or concave surface.
A somewhat different method of observation was next followed: Fig. 3. Brassica oleracea: circumnutating movement of buried and arched hypocotyl (dimly illuminated from above), traced on horizontal glass during 45 hours. Movement of bead of filament magnified about 25 times, and here reduced to one-half of original scale.
as soon as the earth with seeds in a pot began to crack, the surface was removed in parts to the depth of .2 inch; and a filament was fixed to the basal leg of a buried and arched hypocotyl, just above the summit of the radicle. The cotyledons were still almost completely enclosed within the much-cracked seed-coats; and these were again covered up with damp adhesive soil pressed pretty firmly down. The movement of the filament was traced (Fig. 3) from 11 A.M. Feb. 5th till 8 A.M. Feb. 7th. By this latter period the cotyledons had been dragged from beneath the pressed-down earth, but the upper part of the hypocotyl still formed nearly a right angle with the lower part. The tracing shows that the arched hypocotyl tends at this early [page 14] age to circumnutate irregularly. On the first day the greater movement (from right to left in the figure) was not in the plane of the vertical and arched hypocotyl, but at right angles to it, or in the plane of the two cotyledons, which were still in close contact. The basal leg of the arch at the time when the filament was affixed to it, was already bowed considerably backwards, or from the cotyledons; had the filament been affixed before this bowing occurred, the chief movement would have been at right angles to that shown in the figure. A filament was attached to another buried hypocotyl of the same age, and it moved in a similar general manner, but the line traced was not so complex. This hypocotyl became almost straight, and the cotyledons were dragged from beneath the ground on the evening of the second day.
Fig. 4. Brassica oleracea: circumnutating movement of buried and arched hypocotyl, with the two legs of the arch tied together, traced on horizontal glass during 33 hours. Movement of the bead of filament magnified about 26 times, and here reduced to one-half original scale.
Before the above observations were made, some arched hypocotyls buried at the depth of a quarter of an inch were uncovered; and in order to prevent the two legs of the arch from beginning to separate at once, they were tied together with fine silk. This was done partly because we wished to ascertain how long the hypocotyl, in its arched condition, would continue to move, and whether the movement when not masked and disturbed by the straightening process, indicated circumnutation. Firstly a filament was fixed to the basal leg of an arched hypocotyl close above the summit of the radicle. The cotyledons were still partially enclosed within the seed-coats. The movement was traced (Fig. 4) from 9.20 A.M. on Dec. [page 15] 23rd to 6.45 A.M. on Dec. 25th. No doubt the natural movement was much disturbed by the two legs having been tied together; but we see that it was distinctly zigzag, first in one direction and then in an almost opposite one. After 3 P.M. on the 24th the arched hypocotyl sometimes remained stationary for a considerable time, and when moving, moved far slower than before. Therefore, on the morning of the 25th, the glass filament was removed from the base of the basal leg, and was fixed horizontally on the summit of the arch, which, from the legs having been tied, had grown broad and almost flat. The movement was now traced during 23 hours (Fig. 5), and we
Fig. 5. Brassica oleracea: circumnutating movement of the crown of a buried and arched hypocotyl, with the two legs tied together, traced on a horizontal glass during 23 hours. Movement of the bead of the filament magnified about 58 times, and here reduced to one-half original scale.
see that the course was still zigzag, which indicates a tendency to circumnutation. The base of the basal leg by this time had almost completely ceased to move.
As soon as the cotyledons have been naturally dragged from beneath the ground, and the hypocotyl has straightened itself by growth along the inner or concave surface, there is nothing to interfere with the free movements of the parts; and the circumnutation now becomes much more regular and clearly displayed, as shown in the following cases:—A seedling was placed in front and near a north-east window with a line joining the [page 16] two cotyledons parallel to the window. It was thus left the whole day so as to accommodate itself to the light. On the following morning a filament was fixed to the midrib of the larger and taller cotyledon (which enfolds the other and smaller one, whilst still within the seed), and a mark being placed close behind, the movement of the whole plant, that is, of the hypocotyl and cotyledon, was traced greatly magnified on a vertical glass. At first the plant bent so much towards the light that it was useless to attempt to trace the movement; but at 10 A.M. heliotropism almost wholly ceased and the first dot was
Fig. 6. Brassica oleracea: conjoint circumnutation of the hypocotyl and cotyledons during 10 hours 45 minutes. Figure here reduced to one-half original scale.
made on the glass. The last was made at 8.45 P.M.; seventeen dots being altogether made in this interval of 10 h. 45 m. (see Fig. 6). It should be noticed that when I looked shortly after 4 P.M. the bead was pointing off the glass, but it came on again at 5.30 P.M., and the course during this interval of 1 h. 30 m. has been filled up by imagination, but cannot be far from correct. The bead moved seven times from side to side, and thus described 3 ellipses in 10 3/4 h.; each being completed on an average in 3 h. 4 m.
On the previous day another seedling had been observed under similar conditions, excepting that the plant was so [page 17] placed that a line joining the two cotyledons pointed towards the window; and the filament was attached to the smaller cotyledon on the side furthest from the window. Moreover the plant was now for the first time placed in this position. The cotyledons bowed themselves greatly towards the light from 8 to 10.50 A.M., when the first dot was made (Fig. 7). During the
Fig. 7. Brassica oleracea: conjoint circumnutation of the hypocotyl and cotyledons, from 10.50 A.M. to 8 A.M. on the following morning. Tracing made on a vertical glass.
next 12 hours the bead swept obliquely up and down 8 times and described 4 figures representing ellipses; so that it travelled at nearly the same rate as in the previous case. during the night it moved upwards, owing to the sleep-movement of the cotyledons, and continued to move in the same direction till 9 A.M. on the following morning; but this latter movement would not have occurred with seedlings under their natural conditions fully exposed to the light.
By 9.25 A.M. on this second day the same cotyledon had [page 18] begun to fall, and a dot was made on a fresh glass. The movement was traced until 5.30 P.M. as shown in (Fig. 8), which is given, because the course followed was much more irregular than on the two previous occasions. During these 8 hours the bead changed its course greatly 10 times. The upward movement of the cotyledon during the afternoon and early part of the night is here plainly shown.
Fig. 8. Brassica oleracea: conjoint circumnutation of the hypocotyl and cotyledons during 8 hours. Figure here reduced to one-third of the original scale, as traced on a vertical glass.
As the filaments were fixed in the three last cases to one of the cotyledons, and as the hypocotyl was left free, the tracings show the movement of both organs conjoined; and we now wished to ascertain whether both circumnutated. Filaments were therefore fixed horizontally to two hypocotyls close beneath the petioles of their cotyledons. These seedlings had stood for two days in the same position before a north-east window. In the morning, up to about 11 A.M., they moved in zigzag lines towards the light; and at night they again became almost upright through apogeotropism. After about 11 A.M. they moved a little back from the light, often crossing and recrossing their former path in zigzag lines. the sky on this day varied much in brightness, and these observations merely proved that the hypocotyls were continually moving in a manner resembling circumnutation. On a previous day which was uniformly cloudy, a hypocotyl was firmly secured to a little stick, and a filament was fixed to the larger of the two cotyledons, and its movement was traced on a vertical glass. It fell greatly from 8.52 A.M., when the first dot was made, till 10.55 A.M.; it then rose greatly until 12.17 P.M. Afterwards it fell a little and made a loop, but by 2.22 P.M. it had risen a little and continued rising till 9.23 P.M., when it made another loop, and at 10.30 P.M. was again rising. These observations show that the cotyledons move [page 19] vertically up and down all day long, and as there was some slight lateral movement, they circumnutated.
Fig. 9. Brassica oleracea: circumnutation of hypocotyl, in darkness, traced on a horizontal glass, by means of a filament with a bead fixed across its summit, between 9.15 A.M. and 8.30 A.M. on the following morning. Figure here reduced to one-half of original scale.
The cabbage was one of the first plants, the seedlings of which were observed by us, and we did not then know how far the circumnutation of the different parts was affected by light. Young seedlings were therefore kept in complete darkness except for a minute or two during each observation, when they were illuminated by a small wax taper held almost vertically above them. During the first day the hypocotyl of one changed its course 13 times (see Fig. 9); and it deserves notice that the longer axes of the figures described often cross one another at right or nearly right angles. Another seedling was observed in the same manner, but it was much older, for it had formed a true leaf a quarter of an inch in length, and the hypocotyl was 1 3/8 inch in height. The figure traced was a very complex one, though the movement was not so great in extent as in the last case.
The hypocotyl of another seedling of the same age was secured to a little stick, and a filament having been fixed to the midrib of one of the cotyledons, the movement of the bead was traced during 14 h. 15 m. (see Fig. 10) in darkness. It should be noted that the chief movement of the cotyledons, namely, up and down, would be shown on a horizontal glass-plate only by the lines in the direction of the midrib (that is, [page 20] up and down, as Fig. 10 here stands) being a little lengthened or shortened; whereas any lateral movement would be well exhibited. The present tracing shows that the cotyledon did thus move laterally (that is, from side to side in the tracing) 12 times in the 14 h. 15 m. of observation. Therefore the cotyledons certainly circumnutated, though the chief movement was up and down in a vertical plane.
Fig 10. Brassica oleracea: circumnutation of a cotyledon, the hypocotyl having been secured to a stick, traced on a horizontal glass, in darkness, from 8.15 A.M. to 10.30 P.M. Movement of the bead of the filament magnified 13 times.
Rate of Movement.—The movements of the hypocotyls and cotyledons of seedling cabbages of different ages have now been sufficiently illustrated. With respect to the rate, seedlings were placed under the microscope with the stage removed, and with a micrometer eye-piece so adjusted that each division equalled 1/500 inch; the plants were illuminated by light passing through a solution of bichromate of potassium so as to eliminate heliotropism. Under these circumstances it was interesting to observe how rapidly the circumnutating apex of a cotyledon passed across the divisions of the micrometer. Whilst travelling in any direction the apex generally oscillated backwards and forwards to the extent of 1/500 and sometimes of nearly 1/250 of an inch. These oscillations were quite different from the trembling caused by any disturbance in the same room or by the shutting of a distant door. The first seedling observed was nearly two inches in height and had been etiolated by having been grown in darkness. The tip of the cotyledon passed across 10 divisions of the micrometer, that is, 1/50 of an inch, in 6 m. 40 s. Short glass filaments were then fixed vertically to the hypocotyls of several seedlings so as to project a little above the cotyledons, thus exaggerating the rate of movement; but only a few of the observations thus made are worth giving. The most remarkable fact was the oscillatory movement above described, and the difference of rate at which the point crossed the divisions of the micrometer, after short intervals of time. For instance, a tall not-etiolated seedling had been kept for 14 h. in darkness; it was exposed before a north-east window for only [page 21] two or three minutes whilst a glass filament was fixed vertically to the hypocotyl; it was then again placed in darkness for half an hour and afterwards observed by light passing through bichromate of potassium. The point, oscillating as usual, crossed five divisions of the micrometer (i.e. 1/100 inch) in 1 m. 30 s. The seedling was then left in darkness for an hour, and now it required 3 m. 6 s. to cross one division, that is, 15 m. 30 s. to have crossed five divisions. Another seedling, after being occasionally observed in the back part of a northern room with a very dull light, and left in complete darkness for intervals of half an hour, crossed five divisions in 5 m. in the direction of the window, so that we concluded that the movement was heliotropic. But this was probably not the case, for it was placed close to a north-east window and left there for 25 m., after which time, instead of moving still more quickly towards the light, as might have been expected, it travelled only at the rate of 12 m. 30 s. for five divisions. It was then again left in complete darkness for 1 h., and the point now travelled in the same direction as before, but at the rate of 3 m. 18 s. for five divisions.
We shall have to recur to the cotyledons of the cabbage in a future chapter, when we treat of their sleep-movements. The circumnutation, also, of the leaves of fully-developed plants will hereafter be described.
Fig. 11. Githago segetum: circumnutation of hypocotyl, traced on a horizontal glass, by means of a filament fixed transversely across its summit, from 8.15 A.M. to 12.15 P.M. on the following day. Movement of bead of filament magnified about 13 times, here reduced to one-half the original scale.
Githago segetum (Caryophylleae).—A young seedling was dimly illuminated from above, and the circumnutation of the hypo- [page 22] cotyl was observed during 28 h., as shown in Fig. 11. It moved in all directions; the lines from right and to left in the figure being parallel to the blades of the cotyledons. The actual distance travelled from side to side by the summit of the hypocotyl was about .2 of an inch; but it was impossible to be accurate on this head, as the more obliquely the plant was viewed, after it had moved for some time, the more the distances were exaggerated.
We endeavoured to observe the circumnutation of the cotyledons, but as they close together unless kept exposed to a moderately bright light, and as the hypocotyl is extremely heliotropic, the necessary arrangements were too troublesome. We shall recur to the nocturnal or sleep-movements of the cotyledons in a future chapter.
Fig. 12. Gossypium: circumnutation of hypocotyl, traced on a horizontal glass, from 10.30 A.M. to 9.30 A.M. on following morning, by means of a filament fixed across its summit. Movement of bead of filament magnified about twice; seedling illuminated from above.
Gossypium (var. Nankin cotton) (Malvaceae).—The circumnutation of a hypocotyl was observed in the hot-house, but the movement was so much exaggerated that the bead twice passed for a time out of view. It was, however, manifest that two somewhat irregular ellipses were nearly completed in 9 h. Another seedling, 1 in. in height, was then observed during 23 h.; but the observations were not made at sufficiently short intervals, as shown by the few dots in Fig. 12, and the tracing was not now sufficiently enlarged. Nevertheless there could be no doubt about the circumnutation of the hypocotyl, which described in 12 h. a figure representing three irregular ellipses of unequal sizes.
The cotyledons are in constant movement up and down during the whole day, and as they offer the unusual case of moving downwards late in the evening and in the early part of the night, many observations were made on them. A filament was fixed along the middle of one, and its movement traced on a vertical glass; but the tracing is not given, as the hypocotyl was not secured, so that it was impossible to distinguish clearly between its movement and that of the cotyledon. The cotyledons rose from 10.30 A.M. to about 3 P.M.; they then sank till 10 P.M., rising, however, greatly in the latter part of the night. [page 23] The angles above the horizon at which the cotyledons of another seedling stood at different hours is recorded in the following short table: —
Oct. 20 2.50 P.M...25o above horizon. Oct. 20 4.20 P.M...22o above horizon. Oct. 20 5.20 P.M...15o above horizon. Oct. 20 10.40 P.M...8o above horizon. Oct. 21 8.40 A.M...28o above horizon. Oct. 21 11.15 A.M...35o above horizon. Oct. 21 9.11 P.M...10o below horizon.
The position of the two cotyledons was roughly sketched at various hours with the same general result.
In the following summer, the hypocotyl of a fourth seedling was secured to a little stick, and a glass filament with triangles of paper having been fixed to one of the cotyledons, its movements were traced on a vertical glass under a double skylight in the house. The first dot was made at 4.20 P.M. June 20th; and the cotyledon fell till 10.15 P.M. in a nearly straight line. Just past midnight it was found a little lower and somewhat to one side. By the early morning, at 3.45 A.M., it had risen greatly, but by 6.20 A.M. had fallen a little. During the whole of this day (21st) it fell in a slightly zigzag line, but its normal course was disturbed by the want of sufficient illumination, for during the night it rose only a little, and travelled irregularly during the whole of the following day and night of June 22nd. The ascending and descending lines traced during the three days did not coincide, so that the movement was one of circumnutation. This seedling was then taken back to the hot-house, and after five days was inspected at 10 P.M., when the cotyledons were found hanging so nearly vertically down, that they might justly be said to have been asleep. On the following morning they had resumed their usual horizontal position.
Oxalis rosea (Oxalideae).—The hypocotyl was secured to a little stick, and an extremely thin glass filament, with two triangles of paper, was attached to one of the cotyledons, which was .15 inch in length. In this and the following species the end of the petiole, where united to the blade, is developed into a pulvinus. The apex of the cotyledon stood only 5 inches from the vertical glass, so that its movement was not greatly exaggerated as long as it remained nearly horizontal; but in the course of the day it both rose considerably above and fell beneath a horizontal position, and then of course the movement was much exaggerated. [page 24] In Fig. 13 its course is shown from 6.45 A.M. on June 17th, to 7.40 A.M. on the following morning; and we see that during the daytime, in the course of 11 h. 15 m., it travelled thrice down and twice up. After 5.45 P.M. it moved rapidly downwards, and in an hour or two depended vertically; it thus remained all night asleep. This position could not be represented on the vertical glass nor in the figure here given. By 6.40 A.M. on the following morning (18th) both cotyledons had risen greatly, and they continued to rise until 8 A.M., when they stood almost horizontally. Their movement was traced during the whole of this day and until the next morning; but a tracing is not given, as it was closely similar to Fig. 13, excepting that the lines were more zigzag. The cotyledons moved 7 times, either upwards or downwards; and at about 4 P.M. the great nocturnal sinking movement commenced.
Fig. 13. Oxalis rosea: circumnutation of cotyledons, the hypocotyl being secured to a stick; illuminated from above. Figure here given one-half of original scale.
Another seedling was observed in a similar manner during nearly 24 h., but with the difference that the hypocotyl was left free. The movement also was less magnified. Between 8.12 A.M. and 5 P.M. on the 18th, the apex of the cotyledon moved 7 times upwards or downwards (Fig. 14). The nocturnal sinking movement, which is merely a great increase of one of the diurnal oscillations, commenced about 4 P.M.
Oxalis Valdiviana.—This species is interesting, as the coty- [page 25] ledons rise perpendicularly upwards at night so as to come into close contact, instead of sinking vertically downwards, as in the case of O. rosea. A glass filament was fixed to a cotyledon, .17 of an inch in length, and the hypocotyl was left free. On
Fig. 14. Oxalis rosea: conjoint circumnutation of the cotyledons and hypocotyl, traced from 8.12 A.M. on June 18th to 7.30 A.M. 19th. The apex of the cotyledon stood only 3 3/4 inches from the vertical glass. Figure here given one-half of original scale.
Fig. 15. Oxalis Valdiviana: conjoint circumnutation of a cotyledon and the hypocotyl, traced on vertical glass, during 24 hours. Figure here given one-half of original scale; seedling illuminated from above.
the first day the seedling was placed too far from the vertical glass; so that the tracing was enormously exaggerated and the movement could not be traced when the cotyledon either rose or sank much; but it was clearly seen that the cotyledons rose thrice and fell twice between 8.15 A.M. and 4.15 P.M. Early on the following morning (June 19th) the apex of a cotyledon was [page 26] placed only 1 7/8 inch from the vertical glass. At 6.40 A.M. it stood horizontally; it then fell till 8.35, and then rose. Altogether in the course of 12 h. it rose thrice and fell thrice, as may be seen in Fig. 15. The great nocturnal rise of the cotyledons usually commences about 4 or 5 P.M., and on the following morning they are expanded or stand horizontally at about 6.30 A.M. In the present instance, however, the great nocturnal rise did not commence till 7 P.M.; but this was due to the hypocotyl having from some unknown cause temporarily bent to the left side, as is shown in the tracing. To ascertain positively that the hypocotyl circumnutated, a mark was placed at 8.15 P.M. behind the two now closed and vertical cotyledons; and the movement of a glass filament fixed upright to the top of the hypocotyl was traced until 10.40 P.M. During this time it moved from side to side, as well as backwards and forwards, plainly showing circumnutation; but the movement was small in extent. Therefore Fig. 15 represents fairly well the movements of the cotyledons alone, with the exception of the one great afternoon curvature to the left.
Oxalis corniculata (var. cuprea).—The cotyledons rise at night to a variable degree above the horizon, generally about 45o: those on some seedlings between 2 and 5 days old were found to be in continued movement all day long; but the movements were more simple than in the last two species. This may have partly resulted from their not being sufficiently illuminated whilst being observed, as was shown by their not beginning to rise until very late in the evening.
Oxalis (Biophytum) sensitiva.—The cotyledons are highly remarkable from the amplitude and rapidity of their movements during the day. The angles at which they stood above or beneath the horizon were measured at short intervals of time; and we regret that their course was not traced during the whole day. We will give only a few of the measurements, which were made whilst the seedlings were exposed to a temperature of 22 1/2o to 24 decrees C. One cotyledon rose 70o in 11 m.; another, on a distinct seedling, fell 80o in 12 m. Immediately before this latter fall the same cotyledon had risen from a vertically downward to a vertically upward position in 1 h. 48 m., and had therefore passed through 180o in under 2 h. We have met with no other instance of a circumnutating movement of such great amplitude as 180o; nor of such rapidity of movement as the passage through 80o in 12 m. The cotyledons of this plant sleep at night by rising [page 27] vertically and coming into close contact. This upward movement differs from one of the great diurnal oscillations above described only by the position being permanent during the night and by its periodicity, as it always commences late in the evening.
Tropaeolum minus (?) (var. Tom Thumb) (Tropaeoleae).—The cotyledons are hypogean, or never rise above the ground. By removing the soil a buried epicotyl or plumule was found, with its summit arched abruptly downwards, like the arched hypocotyl of the cabbage previously described. A glass filament with a bead at its end was affixed to the basal half or leg, just above the hypogean cotyledons, which were again almost surrounded by loose earth. The tracing (Fig. 16) shows the course of the bead during 11 h. After the last dot given in the figure, the bead moved to a great distance, and finally off the glass, in the direction indicated by the broken line. This great movement, due to increased growth along the concave surface of the arch, was caused by the basal leg bending backwards from the upper part, that is in a direction opposite to the dependent tip, in the same manner as occurred with the hypocotyl of the cabbage. Another buried and arched epicotyl was observed in the same manner, excepting that the two legs of the arch were tied together with fine silk for the sake of preventing the great movement just mentioned. It moved, however, in the evening in the same direction as before, but the line followed was not so straight. During the morning the tied arch moved in an irregularly circular, strongly zigzag course, and to a greater distance than in the previous case, as was shown in a tracing, magnified 18 times. The movements of a young plant bearing a few leaves and of a mature plant, will hereafter be described.
Fig. 16. Tropaeolum minus (?): circumnutation of buried and arched epicotyl, traced on a horizontal glass, from 9.20 A.M. to 8.15 P.M. Movement of bead of filament magnified 27 times. [page 28]
Citrus aurantium (Orange) (Aurantiaceae).—The cotyledons are hypogean. The circumnutation of an epicotyl, which at the close of our observations was .59 of an inch (15 mm.) in height above the ground, is shown in the annexed figure (Fig. 17), as observed during a period of 44 h. 40 m.
Fig. 17. Citrus aurantium: circumnutation of epicotyl with a filament fixed transversely near its apex, traced on a horizontal glass, from 12.13 P.M. on Feb. 20th to 8.55 A.M. on 22nd. The movement of the bead of the filament was at first magnified 21 times, or 10 1/2, in figure here given, and afterwards 36 times, or 18 as here given; seedling illuminated from above.
Aesculus hippocastanum (Hippocastaneae).—Germinating seeds were placed in a tin box, kept moist internally, with a sloping bank of damp argillaceous sand, on which four smoked glass-plates rested, inclined at angles of 70o and 65o with the horizon. The tips of the radicles were placed so as just to touch the upper end of the glass-plates, and, as they grew downwards they pressed lightly, owing to geotropism, on the smoked surfaces, and left tracks of their course. In the middle part of each track the glass was swept clean, but the margins were much blurred and irregular. Copies of two of these tracks (all four being nearly alike) were made on tracing paper placed over the glass-plates after they had been varnished; and they are as exact as possible considering the nature of the margins (Fig. 18). They suffice to show that there was some lateral, almost serpentine movement, and that the tips in their downward course pressed with unequal force on the plates, as [page 29] the tracks varied in breadth. The more perfectly serpentine tracks made by the radicles of Phaseolus multiflorus and Vicia faba (presently to be described), render it almost certain that the radicles of the present plant circumnutated.
Fig. 18. Aesculus hippocastanum: outlines of tracks left on inclined glass-plates by tips of radicles. In A the plate was inclined at 70o with the horizon, and the radicle was 1.9 inch in length, and .23 inch in diameter at base. In B the plate was inclined 65o with the horizon, and the radicle was a trifle larger.
Phaseolus multiflorus (Leguminosae).—Four smoked glass-plates were arranged in the same manner as described under Aesculus, and the tracks left by the tips of four radicles of the present plant, whilst growing downwards, were photographed as transparent objects. Three of them are here exactly copied (Fig. 19). Their serpentine courses show that the tips moved regularly from side to side; they also pressed alternately with greater or less force on the plates, sometimes rising up and leaving them altogether for a very short distance; but this was better seen on the original plates than in the copies. These radicles therefore were continually moving in all directions—that is, they circumnutated. The distance between the extreme right and left positions of the radicle A, in its lateral movement, was 2 mm., as ascertained by measurement with an eye-piece micrometer.
Fig. 19. Phaseolus multiflorus: tracks left on inclined smoked glass-plates by tips of radicles in growing downwards. A and C, plates inclined at 60o, B inclined at 68o with the horizon.
Vicia faba (Common Bean) (Leguminosae).—Radicle.—Some beans were allowed to germinate on bare sand, and after one had protruded its radicle to a length of .2 of an inch, it was turned upside down, so that the radicle, which was kept in damp air, now stood upright. A filament, nearly an inch in length, was affixed obliquely near its tip; and the movement of the terminal bead was traced from 8.30 A.M. to 10.30 P.M., as shown in Fig. 18. The radicle at first changed its course twice [page 30] abruptly, then made a small loop and then a larger zigzag curve. During the night and till 11 A.M. on the following
Fig. 20. Vicia faba: circumnutation of a radicle, at first pointing vertically upwards, kept in darkness, traced on a horizontal glass, during 14 hours. Movement of bead of filament magnified 23 times, here reduced to one-half of original scale.
morning, the bead moved to a great distance in a nearly straight line, in the direction indicated by the broken line in the figure. This resulted from the tip bending quickly downwards, as it had now become much declined, and had thus gained a position highly favourable for the action of geotropism. Fig. 21. Vicia faba: tracks left on inclined smoked glass-plates, by tips of radicles in growing downwards. Plate C was inclined at 63o, plates A and D at 71o, plate B at 75o, and plate E at a few degrees beneath the horizon. [page 31]
We next experimented on nearly a score of radicles by allowing them to grow downwards over inclined plates of smoked glass, in exactly the same manner as with Aesculus and Phaseolus. Some of the plates were inclined only a few degrees beneath the horizon, but most of them between 60o and 75o. In the latter cases the radicles in growing downwards were deflected only a little from the direction which they had followed whilst germinating in sawdust, and they pressed lightly on the glass-plates (Fig. 21). Five of the most distinct tracks are here copied, and they are all slightly sinuous, showing circumnutation. Moreover, a close examination of almost every one of the tracks clearly showed that the tips in their downward course had alternately pressed with greater or less force on the plates, and had sometimes risen up so as nearly to leave them for short intervals. The distance between the extreme right and left positions of the radicle A was 0.7 mm., ascertained in the same manner as in the case of Phaseolus.
Epicotyl.—At the point where the radicle had protruded from a bean laid on its side, a flattened solid lump projected .1 of an inch, in the same horizontal plane with the bean. This protuberance consisted of the convex summit of the arched epicotyl; and as it became developed the two legs of the arch curved themselves laterally upwards, owing to apogeotropism, at such a rate that the arch stood highly inclined after 14 h., and vertically in 48 h. A filament was fixed to the crown of the protuberance before any arch was visible, but the basal half grew so quickly that on the second morning the end of the filament was bowed greatly downwards. It was therefore removed and fixed lower down. The line traced during these two days extended in the same general direction, and was in parts nearly straight, and in others plainly zigzag, thus giving some evidence of circumnutation.
As the arched epicotyl, in whatever position it may be placed, bends quickly upwards through apogeotropism, and as the two legs tend at a very early age to separate from one another, as soon as they are relieved from the pressure of the surrounding earth, it was difficult to ascertain positively whether the epicotyl, whilst remaining arched, circumnutated. Therefore some rather deeply buried beans were uncovered, and the two legs of the arches were tied together, as had been done with the epicotyl of Tropaeolum and the hypocotyl of the Cabbage. The movements of the tied arches were traced in the usual manner on [page 32] two occasions during three days. But the tracings made under such unnatural conditions are not worth giving; and it need only be said that the lines were decidedly zigzag, and that small loops were occasionally formed. We may therefore conclude that the epicotyl circumnutates whilst still arched and before it has grown tall enough to break through the surface of the ground.
In order to observe the movements of the epicotyl at a somewhat more advanced age, a filament was fixed near the base of one which was no longer arched, for its upper half now formed a right angle with the lower half. This bean had germinated on bare damp sand, and the epicotyl began to straighten itself much sooner than would have occurred if it had been properly planted. The course pursued during 50 h. (from 9 A.M. Dec. 26th, to 11 A.M. 28th) is here shown (Fig. 22); and we see Fig. 22. Vicia faba: circumnutation of young epicotyl, traced in darkness during 50 hours on a horizontal glass. Movement of bead of filament magnified 20 times, here reduced to one-half of original scale.
that the epicotyl circumnutated during the whole time. Its basal part grew so much during the 50 h. that the filament at the end of our observations was attached at the height of .4 inch above the upper surface of the bean, instead of close to it. If the bean had been properly planted, this part of the epicotyl would still have been beneath the soil.
Late in the evening of the 28th, some hours after the above observations were completed, the epicotyl had grown much straighter, for the upper part now formed a widely open angle with the lower part. A filament was fixed to the upright basal part, higher up than before, close beneath the lowest scale-like process or homologue of a leaf; and its movement was traced [page 33] during 38 h. (Fig. 23). We here again have plain evidence of continued circumnutation. Had the bean been properly planted, the part of the epicotyl to which the filament was attached, the
Fig. 23. Vicia faba: circumnutation of the same epicotyl as in Fig. 22, a little more advanced in age, traced under similar conditions as before, from 8.40 A.M. Dec. 28th, to 10.50 A.M. 30th. Movement of bead here magnified 20 times.
movement of which is here shown, would probably have just risen above the surface of the ground.
Lathyrus nissolia (Leguminosae).—This plant was selected for observation from being an abnormal form with grass-like leaves.
Fig. 24. Lathyrus nissolia: circumnutation of stem of young seedling, traced in darkness on a horizontal glass, from 6.45 A.M. Nov. 22nd, to 7 A.M. 23rd. Movement of end of leaf magnified about 12 times, here reduced to one-half of original scale.
The cotyledons are hypogean, and the epicotyl breaks through the ground in an arched form. The movements of a stem, 1.2 inch in height, consisting of three internodes, the lower one almost wholly subterranean, and the upper one bearing a short, [page 34] narrow leaf, is shown during 24 h., in Fig. 24. No glass filament was employed, but a mark was placed beneath the apex of the leaf. The actual length of the longer of the two ellipses described by the stem was about .14 of an inch. On the previous day the chief line of movement was nearly at right angles to that shown in the present figure, and it was more simple.
Cassia tora* (Leguminosae).—A seedling was placed before a
Fig. 25. Cassia tora: conjoint circumnutation of cotyledons and hypocotyl, traced on vertical glass, from 7.10 A.M. Sept. 25th to 7.30 A.M. 26th. Figure here given reduced to one-half of original scale.
* Seeds of this plant, which grew near the sea-side, were sent to us by Fritz Mller from S. Brazil. The seedlings did not flourish or flower well with us; they were sent to Kew, and were pronounced not to be distinguishable from C. tora. [page 35]
north-east window; it bent very little towards it, as the hypocotyl which was left free was rather old, and therefore not highly heliotropic. A filament had been fixed to the midrib of one of the cotyledons, and the movement of the whole seedling was traced during two days. The circumnutation of the hypocotyl is quite insignificant compared with that of the cotyledons. These rise up vertically at night and come into close contact; so that they may be said to sleep. This seedling was so old that a very small true leaf had been developed, which at night was completely hidden by the closed cotyledons. On Sept. 24th, between 8 A.M. and 5 P.M., the cotyledons moved five times up and five times down; they therefore described five irregular ellipses in the course of the 9 h. The great nocturnal rise commenced about 4.30 P.M.
On the following morning (Sept. 25th) the movement of the same cotyledon was again traced in the same manner during 24 h.; and a copy of the tracing is here given (Fig. 25). The morning was cold, and the window had been accidentally left open for a short time, which must have chilled the plant; and this probably prevented it from moving quite as freely as on the previous day; for it rose only four and sank only four times during the day, one of the oscillations being very small. At 7.10 A.M., when the first dot was made, the cotyledons were not fully open or awake; they continued to open till about 9 A.M., by which time they had sunk a little beneath the horizon: by 9.30 A.M. they had risen, and then they oscillated up and down; but the upward and downward lines never quite coincided. At about 4.30 P.M. the great nocturnal rise commenced. At 7 A.M. on the following morning (Sept. 26th) they occupied nearly the same level as on the previous morning, as shown in the diagram: they then began to open or sink in the usual manner. The diagram leads to the belief that the great periodical daily rise and fall does not differ essentially, excepting in amplitude, from the oscillations during the middle of the day.
Lotus Jacoboeus (Leguminosae).—The cotyledons of this plant, after the few first days of their life, rise so as to stand almost, though rarely quite, vertically at night. They continue to act in this manner for a long time even after the development of some of the true leaves. With seedlings, 3 inches in height, and bearing five or six leaves, they rose at night about 45o. They continued to act thus for about an additional fortnight. Subsequently they remained horizontal at night, though still green [page 36] and at last dropped off. Their rising at night so as to stand almost vertically appears to depend largely on temperature; for when the seedlings were kept in a cool house, though they still continued to grow, the cotyledons did not become vertical at night. It is remarkable that the cotyledons do not generally rise at night to any conspicuous extent during the first four or five days after germination; but the period was extremely variable with seedlings kept under the same conditions; and many were observed. Glass filaments with minute triangles of paper were fixed to the cotyledons (1 mm. in breadth) of two seedlings, only 24 h. old, and the hypocotyl was secured to a stick; their movements greatly magnified were traced, and they certainly circumnutated the whole time on a small scale, but they did not exhibit any distinct nocturnal and diurnal movement. The hypocotyls, when left free, circumnutated over a large space.
Another and much older seedling, bearing a half-developed leaf, had its movements traced in a similar manner during the three first days and nights of June; but seedlings at this age appear to be very sensitive to a deficiency of light; they were observed under a rather dim skylight, at a temperature of between 16o to 17 1/2o C.' and apparently, in consequence of these conditions, the great daily movement of the cotyledons ceased on the third day. During the first two days they began rising in the early afternoon in a nearly straight line, until between 6 and 7 P.M., when they stood vertically. During the latter part of the night, or more probably in the early morning, they began to fall or open, so that by 6.45 A.M. they stood fully expanded and horizontal. They continued to fall slowly for some time, and during the second day described a single small ellipse, between 9 A.M. and 2 P.M., in addition to the great diurnal movement. The course pursued during the whole 24 h. was far less complex than in the foregoing case of Cassia. On the third morning they fell very much, and then circumnutated on a small scale round the same spot; by 8.20 P.M. they showed no tendency to rise at night. Nor did the cotyledons of any of the many other seedlings in the same pot rise; and so it was on the following night of June 5th. The pot was then taken back into the hot-house, where it was exposed to the sun, and on the succeeding night all the cotyledons rose again to a high angle, but did not stand quite vertically. On each of the above days the line representing the great nocturnal [page 37] rise did not coincide with that of the great diurnal fall, so that narrow ellipses were described, as is the usual rule with circumnutating organs. The cotyledons are provided with a pulvinus, and its development will hereafter be described.
Mimosa pudica (Leguminosae).—The cotyledons rise up vertically at night, so as to close together. Two seedlings were observed in the greenhouse (temp. 16o to 17o C. or 63o to 65o F.). Their hypocotyls were secured to sticks, and glass filaments bearing little triangles of paper were affixed to the cotyledons of both. Their movements were traced on a vertical glass during 24 h. on November 13th. The pot had stood for some time in the same position, and they were chiefly illuminated through the glass-roof. The cotyledons of one of these seedlings moved downward in the morning till 11.30 A.M., and then rose, moving rapidly in the evening until they stood vertically, so that in this case there was simply a single great daily fall and rise. The other seedling behaved rather differently, for it fell in the morning until 11.30 A.M., and then rose, but after 12.10 P.M. again fell; and the great evening rise did not begin until 1.22 P.M. On the following morning this cotyledon had fallen greatly from its vertical position by 8.15 A.M. Two other seedlings (one seven and the other eight days old) had been previously observed under unfavourable circumstances, for they had been brought into a room and placed before a north-east window, where the temperature was between only 56o and 57o F. They had, moreover, to be protected from lateral light, and perhaps were not sufficiently illuminated. Under these circumstances the cotyledons moved simply downwards from 7 A.M. till 2 P.M., after which hour and during a large part of the night they continued to rise. Between 7 and 8 A.M. on the following morning they fell again; but on this second and likewise on the third day the movements became irregular, and between 3 and 10.30 P.M. they circumnutated to a small extent about the same spot; but they did not rise at night. Nevertheless, on the following night they rose as usual.
Cytisus fragrans (Leguminosae).—Only a few observations were made on this plant. The hypocotyl circumnutated to a considerable extent, but in a simple manner—namely, for two hours in one direction, and then much more slowly back again in a zigzag course, almost parallel to the first line, and beyond the starting-point. It moved in the same direction all night, but next morning began to return. The cotyledons continually [page 38] move both up and down and laterally; but they do not rise up at night in a conspicuous manner.
Lupinus luteus (Leguminosae).—Seedlings of this plant were observed because the cotyledons are so thick (about .08 of an inch) that it seemed unlikely that they would move. Our observations were not very successful, as the seedlings are strongly heliotropic, and their circumnutation could not be accurately observed near a north-east window, although they had been kept during the previous day in the same position. A seedling was then placed in darkness with the hypocotyl secured to a stick; both cotyledons rose a little at first, and then fell during the rest of the day; in the evening between 5 and 6 P.M. they moved very slowly; during the night one continued to fall and the other rose, though only a little. The tracing was not much magnified, and as the lines were plainly zigzag, the cotyledons must have moved a little laterally, that is, they must have circumnutated.
The hypocotyl is rather thick, about .12 of inch; nevertheless it circumnutated in a complex course, though to a small extent. The movement of an old seedling with two true leaves partially developed, was observed in the dark. As the movement was magnified about 100 times it is not trustworthy and is not given; but there could be no doubt that the hypocotyl moved in all directions during the day, changing its course 19 times. The extreme actual distance from side to side through which the upper part of the hypocotyl passed in the course of 14 hours was only 1/60 of an inch; it sometimes travelled at the rate of 1/50 of an inch in an hour.
Cucurbita ovifera (Cucurbitaceae).—Radicle: a seed which had
Fig. 26. Cucurbita ovifera: course followed by a radicle in bending geotropically downwards, traced on a horizontal glass, between 11.25 A.M. and 10.25 P.M.; the direction during the night is indicated by the broken line. Movement of bead magnified 14 times.
germinated on damp sand was fixed so that the slightly curved radicle, which was only .07 inch in length, stood almost vertically [page 39] upwards, in which position geotropism would act at first with little power. A filament was attached near to its base, and projected at about an angle of 45o above the horizon. The general course followed during the 11 hours of observation and during the following night is shown in the accompanying diagram (Fig. 26), and was plainly due to geotropism; but it was also clear that the radicle circumnutated. By the next morning the tip had curved so much downwards that the filament, instead of projecting at 45o above the horizon, was nearly horizontal. Another germinating seed was turned upside down and covered with damp sand; and a filament was fastened to the radicle so as to project at an angle of about 50o above the horizon; this radicle was .35 of an inch in length and a little curved. The course pursued was mainly governed, as in the last case, by geotropism, but the line traced during 12 hours and magnified as before was more strongly zigzag, again showing circumnutation.
Four radicles were allowed to grow downwards over plates of smoked glass, inclined at 70o to the horizon, under the
Fig. 27. Cucurbita ovifera: tracks left by tips of radicles in growing downwards over smoked glass-plates, inclined at 70o to the horizon.
Fig. 28. Cucurbita ovifera: circumnutation of arched hypocotyl at a very early age, traced in darkness on a horizontal glass, from 8 A.M. to 10.20 A.M. on the following day. The movement of the bead magnified 20 times, here reduced to one-half of original scale.
same conditions as in the cases of Aesculus, Phaseolus, and Vicia. Facsimiles are here given (Fig. 27) of two of these tracks; and a third short one was almost as plainly serpentine as that at A. It was also manifest by a greater or less amount of soot having been swept off the glasses, that the tips had [page 40] pressed alternately with greater and less force on them. There must, therefore, have been movement in at least two planes at right angles to one another. These radicles were so delicate that they rarely had the power to sweep the glasses quite clean. One of them had developed some lateral or secondary rootlets, which projected a few degrees beneath the horizon; and it is an important fact that three of them left distinctly serpentine tracks on the smoked surface, showing beyond doubt that they had circumnutated like the main or primary radicle. But the tracks were so slight that they could not be traced and copied after the smoked surface had been varnished.