and causes the upper part to bend towards its source.
The tips of 15 radicles were cauterised with nitrate of silver, and they grew as well as those above described with greased tips. After an interval of 24 h., 9 of them were not at all curved towards the bottom of the sieve; 2 were curved towards it at angles of 20° and 12° from their former vertical position, and 4 had come into close contact with it. Thus the destruction of the tip for a length of about 1 mm.
prevented the curvature of the greater number of these radicles to the adjoining damp surface. Of 24 control specimens, 23 were bent to the sieve, and on a second occasion 15 out of 16 were similarly curved in a greater or less degree. These control trials are included in those given in the foregoing paragraph.
Avena sativa.—The tips of 13 radicles, which projected between 2 and 4 mm. from the bottom of the sieve, many of them not quite perpendicularly downwards, were coated with the black grease for a length of from 1 to 1½ mm. The sieves were inclined at 30° with the horizon. The greater number of these radicles were examined after 22 h., and a few after 25 h., and within these intervals they had grown so quickly as to have nearly doubled their lengths. With the ungreased radicles the chief seat of curvature is at a distance of not less than between 3.5 and 5.5 mm., and not more than between 7 and 10 mm. from the apex. Out of the 13 radicles with greased tips, 4 had not moved at all towards the sieve; 6 were deflected towards it and from the perpendicular by angles varying between 10° and 35°; and 3 had come into close contact with it. It appears, therefore, at first sight that greasing the tips of these radicles had checked but little their bending to the adjoining damp surface. But the inspection of the sieves on two occasions produced a widely different impression on the mind; for it was impossible to behold the radicles with the black greased tips projecting from the bottom, and all those with ungreased tips, at least 40 to 50 in number, clinging closely to it, and feel any doubt that the greasing had produced a great effect. On close examination only a single ungreased radicle could be found which had not become curved towards the sieve. It is probable that if the tips had been protected by grease for a length of 2 mm. instead of from 1 to 1½ mm., they would not have been affected by the moist air and none would have become curved.
Triticum vulgare.—Analogous trials were made on 8 radicles of the common wheat; and greasing their tips produced much less effect than in the case of the oats. After 22 h., 5 of them had come into contact with the bottom of the sieve; 2 had moved towards it 10° and 15°, and one alone remained perpendicular. Not one of the very numerous ungreased radicles failed to come into close contact with the sieve. These trials were made on Nov. 28th, when the temperature was only 4.8° C. at 10 A.M. We should hardly have thought this case worth notice, had it not been for the following circumstance. In the beginning of October, when the temperature was considerably higher, viz., 12° to 13° C., we found that only a few of the ungreased radicles became bent towards the sieve; and this indicates that sensitiveness to moisture in the air is increased by a low temperature, as we have seen with the radicles of Vicia faba relatively to objects attached to their tips. But in the present instance it is possible that a difference in the dryness of the air may have caused the difference in the results at the two periods.
Finally, the facts just given with respect to Phaseolus multiflorus, Vicia faba, and Avena sativa show, as it seems to us, that a layer of grease spread for a length of 1½ to 2 mm. over the tip of the radicle, or the destruction of the tip by caustic, greatly lessens or quite annuls in the upper and exposed part the power of bending towards a neighbouring source of moisture. We should bear in mind that the part which bends most, lies at some little distance above the greased or cauterised tip; and that the rapid growth of this part, proves that it has not been injured by the tips having been thus treated. In those cases in which the radicles with greased tips became curved, it is possible that the layer of grease was not sufficiently thick wholly to exclude moisture, or that a sufficient length was not thus protected, or, in the case of the caustic, not destroyed. When radicles with greased tips are left to grow for several days in damp air, the grease is drawn out into the finest reticulated threads and dots, with narrow portions of the surface left clean. Such portions would, it is probable, be able to absorb moisture, and thus we can account for several of the radicles with greased tips having become curved towards the sieve after an interval of one or two days. On the whole, we may infer that sensitiveness to a difference in the amount of moisture in the air on the two sides of a radicle resides in the tip, which transmits some influence to the upper part, causing it to bend towards the source of moisture. Consequently, the movement is the reverse of that caused by objects attached to one side of the tip, or by a thin slice being cut off, or by being slightly cauterised. In a future chapter it will be shown that sensitiveness to the attraction of gravity likewise resides in the tip; so that it is the tip which excites the adjoining parts of a horizontally extended radicle to bend towards the centre of the earth.
SECONDARY RADICLES BECOMING VERTICALLY GEOTROPIC BY THE DESTRUCTION OR INJURY OF THE TERMINAL PART OF THE PRIMARY RADICLE.
Sachs has shown that the lateral or secondary radicles of the bean, and probably of other plants, are acted on by geotropism in so peculiar a manner, that they grow out horizontally or a little inclined downwards; and he has further shown[10] the interesting fact, that if the end of the primary radicle be cut off, one of the nearest secondary radicles changes its nature and grows perpendicularly downwards, thus replacing the primary radicle. We repeated this experiment, and planted beans with amputated radicles in friable peat, and saw the result described by Sachs; but generally two or three of the secondary radicles grew perpendicularly downwards. We also modified the experiment, by pinching young radicles a little way above their tips, between the arms of a U-shaped piece of thick leaden wire. The part pinched was thus flattened, and was afterwards prevented from growing thicker. Five radicles had their ends cut off, and served as controls or standards.
Eight were pinched; of these 2 were pinched too severely and their ends died and dropped off; 2 were not pinched enough and were not sensibly affected; the remaining 4 were pinched sufficiently to check the growth of the terminal part, but did not appear otherwise injured. When the U-shaped wires were removed, after an interval of 15 days, the part beneath the wire was found to be very thin and easily broken, whilst the part above was thickened. Now in these four cases, one or more of the secondary radicles, arising from the thickened part just above the wire, had grown perpendicularly downwards. In the best case the primary radicle (the part below the wire being 1½ inch in length) was somewhat distorted, and was not half as long as three adjoining secondary radicles, which had grown vertically, or almost vertically, downwards. Some of these secondary radicles adhered together or had become confluent. We learn from these four cases that it is not necessary, in order that a secondary radicle should assume the nature of a primary one, that the latter should be actually amputated; it is sufficient that the flow of sap into it should be checked, and consequently should be directed into the adjoining secondary radicles; for this seems to be the most obvious result of the primary radicle being pinched between the arms of a U-shaped wire.
This change in the nature of secondary radicles is clearly analogous, as Sachs has remarked, to that which occurs with the shoots of trees, when the leading one is destroyed and is afterwards replaced by one or more of the lateral shoots; for these now grow upright instead of sub-horizontally. But in this latter case the lateral shoots are rendered apogeotropic, whereas with radicles the lateral ones are rendered geotropic. We are naturally led to suspect that the same cause acts with shoots as with roots, namely, an increased flow of sap into the lateral ones. We made some trials with Abies communis and pectinata, by pinching with wire the leading and all the lateral shoots excepting one. But we believe that they were too old when experimented on; and some were pinched too severely, and some not enough. Only one case succeeded, namely, with the spruce-fir.
The leading shoot was not killed, but its growth was checked; at its base there were three lateral shoots in a whorl, two of which were pinched, one being thus killed; the third was left untouched. These lateral shoots, when operated on (July 14th) stood at an angle of 8° above the horizon; by Sept. 8th the unpinched one had risen 35°; by Oct. 4th it had risen 46°, and by Jan. 26th 48°, and it had now become a little curved inwards. Part of this rise of 48° may be attributed to ordinary growth, for the pinched shoot rose 12° within the same period.
It thus follows that the unpinched shoot stood, on Jan. 26th, 56° above the horizon, or 34° from the vertical; and it was thus obviously almost ready to replace the slowly growing, pinched, leading shoot.
Nevertheless, we feel some doubt about this experiment, for we have since observed with spruce-firs growing rather unhealthily, that the lateral shoots near the summit sometimes become highly inclined, whilst the leading shoot remains apparently sound.
A widely different agency not rarely causes shoots which naturally would have brown out horizontally to grow up vertically. The lateral branches of the Silver Fir (A. pectinata) are often affected by a fungus, Æcidium elatinum, which causes the branch to enlarge into an oval knob formed of hard wood, in one of which we counted 24 rings of growth. According to De Bary[11], when the mycelium penetrates a bud beginning to elongate, the shoot developed from it grows vertically upwards. Such upright shoots afterwards produce lateral and horizontal branches; and they then present a curious appearance, as if a young fir-tree had grown out of a ball of clay surrounding the branch. These upright shoots have manifestly changed their nature and become apogeotropic; for if they had not been affected by the Æcidium, they would have grown out horizontally like all the other twigs on the same branches. This change can hardly be due to an increased flow of sap into the part; but the presence of the mycelium will have greatly disturbed its natural constitution.
[11] See his valuable article in ‘Bot. Zeitung,’ 1867, p. 257, on these monstrous growths, which are called in German “Hexenbesen,” or “witch-brooms.”
According to Mr. Meehan,[12] the stems of three species of Euphorbia and of Portulaca oleracea are “normally prostrate or procumbent;” but when they are attacked by an Æcidium, they “assume an erect habit.” Dr.
Stahl informs us that he knows of several analogous cases; and these seem to be closely related to that of the Abies. The rhizomes of Sparganium ramosum grow out horizontally in the soil to a considerable length, or are diageotropic; but F. Elfving found that when they were cultivated in water their tips turned upwards, and they became apogeotropic. The same result followed when the stem of the plant was bent until it cracked or was merely much bowed.[13] [12] ‘Proc. Acad. Nat. Sc. Philadelphia,’ June 16th, 1874, and July 23rd, 1875.
[13] See F. Elfving’s interesting paper in ‘Arbeiten Bot. Institut., in Würzburg,’ vol. ii. 1880, p. 489. Carl Kraus (Triesdorf) had previously observed (‘Flora,’ 1878, p. 324) that the underground shoots of Triticum repens bend vertically up when the parts above ground are removed, and when the rhizomes are kept partly immersed in water.
No explanation has hitherto been attempted of such cases as the foregoing,—namely, of secondary radicles growing vertically downwards, and of lateral shoots growing vertically upwards, after the amputation of the primary radicle or of the leading shoot. The following considerations give us, as we believe, the clue. Firstly, any cause which disturbs the constitution[14] is apt to induce reversion; such as the crossing of two distinct races, or a change of conditions, as when domestic animals become feral. But the case which most concerns us, is the frequent appearance of peloric flowers on the summit of a stem, or in the centre of the inflorescence,—parts which, it is believed, receive the most sap; for when an irregular flower becomes perfectly regular or peloric, this may be attributed, at least partly, to reversion to a primitive and normal type. Even the position of a seed at the end of the capsule sometimes gives to the seedling developed from it a tendency to revert. Secondly, reversions often occur by means of buds, independently of reproduction by seed; so that a bud may revert to the character of a former state many bud-generations ago. In the case of animals, reversions may occur in the individual with advancing age. Thirdly and lastly, radicles when they first protrude from the seed are always geotropic, and plumules or shoots almost always apogeotropic. If then any cause, such as an increased flow of sap or the presence of mycelium, disturbs the constitution of a lateral shoot or of a secondary radicle, it is apt to revert to its primordial state; and it becomes either apogeotropic or geotropic, as the case may be, and consequently grows either vertically upwards or downwards. It is indeed possible, or even probable, that this tendency to reversion may have been increased, as it is manifestly of service to the plant.
[14] The facts on which the following conclusions are founded are given in ‘The Variation of Animals and Plants under Domestication,’ 2nd edit. 1875. On the causes leading to reversion see chap. xii. vol.
ii. and p. 59, chap. xiv. On peloric flowers, chap. xiii. p. 32; and see p. 337 on their position on the plant. With respect to seeds, p.
340. On reversion by means of buds, p. 438, chap. xi. vol. i.
A SUMMARY OF CHAPTER.
A part or organ may be called sensitive, when its irritation excites movement in an adjoining part. Now it has been shown in this chapter, that the tip of the radicle of the bean is in this sense sensitive to the contact of any small object attached to one side by shellac or gum-water; also to a slight touch with dry caustic, and to a thin slice cut off one side. The radicles of the pea were tried with attached objects and caustic, both of which acted. With Phaseolus multiflorus the tip was hardly sensitive to small squares of attached card, but was sensitive to caustic and to slicing. The radicles of Tropaeolum were highly sensitive to contact; and so, as far as we could judge, were those of Gossypium herbaceum, and they were certainly sensitive to caustic. The tips of the radicles of Cucurbita ovifera were likewise highly sensitive to caustic, though only moderately so to contact.
Raphanus sativus offered a somewhat doubtful case. With Æsculus the tips were quite indifferent to bodies attached to them, though sensitive to caustic. Those of Quercus robur and Zea mays were highly sensitive to contact, as were the radicles of the latter to caustic. In several of these cases the difference in sensitiveness of the tip to contact and to caustic was, as we believe, merely apparent; for with Gossypium, Raphanus, and Cucurbita, the tip was so fine and flexible that it was very difficult to attach any object to one of its sides.
With the radicles of Æsculus, the tips were not at all sensitive to small bodies attached to them; but it does not follow from this fact that they would not have been sensitive to somewhat greater continued pressure, if this could have been applied.
The peculiar form of sensitiveness which we are here considering, is confined to the tip of the radicle for a length of from 1 mm. to 1.5 mm. When this part is irritated by contact with any object, by caustic, or by a thin slice being cut off, the upper adjoining part of the radicle, for a length of from 6 or 7 to even 12 mm., is excited to bend away from the side which has been irritated. Some influence must therefore be transmitted from the tip along the radicle for this length. The curvature thus caused is generally symmetrical. The part which bends most apparently coincides with that of the most rapid growth. The tip and the basal part grow very slowly and they bend very little.
Considering the widely separated position in the vegetable series of the several above-named genera, we may conclude that the tips of the radicles of all, or almost all, plants are similarly sensitive, and transmit an influence causing the upper part to bend. With respect to the tips of the secondary radicles, those of Vicia faba, Pisum sativum, and Zea mays were alone observed, and they were found similarly sensitive.
In order that these movements should be properly displayed, it appears necessary that the radicles should grow at their normal rate. If subjected to a high temperature and made to grow rapidly, the tips seem either to lose their sensitiveness, or the upper part to lose the power of bending. So it appears to be if they grow very slowly from not being vigorous, or from being kept at too low a temperature; also when they are forced to germinate in the middle of the winter.
The curvature of the radicle sometimes occurs within from 6 to 8 hours after the tip has been irritated, and almost always within 24 h., excepting in the case of the massive radicles of Æsculus. The curvature often amounts to a rectangle,—that is, the terminal part bends upwards until the tip, which is but little curved, projects almost horizontally. Occasionally the tip, from the continued irritation of the attached object, continues to bend up until it forms a hook with the point directed towards the zenith, or a loop, or even a spire.
After a time the radicle apparently becomes accustomed to the irritation, as occurs in the case of tendrils, for it again grows downwards, although the bit of card or other object may remain attached to the tip. It is evident that a small object attached to the free point of a vertically suspended radicle can offer no mechanical resistance to its growth as a whole, for the object is carried downwards as the radicle elongates, or upwards as the radicle curves upwards. Nor can the growth of the tip itself be mechanically checked by an object attached to it by gum-water, which remains all the time perfectly soft. The weight of the object, though quite insignificant, is opposed to the upward curvature. We may therefore conclude that it is the irritation due to contact which excites the movement. The contact, however, must be prolonged, for the tips of 15 radicles were rubbed for a short time, and this did not cause them to bend. Here then we have a case of specialised sensibility, like that of the glands of Drosera; for these are exquisitely sensitive to the slightest pressure if prolonged, but not to two or three rough touches.
When the tip of a radicle is lightly touched on one side with dry nitrate of silver, the injury caused is very slight, and the adjoining upper part bends away from the cauterised point, with more certainty in most cases than from an object attached on one side. Here it obviously is not the mere touch, but the effect produced by the caustic, which induces the tip to transmit some influence to the adjoining part, causing it to bend away. If one side of the tip is badly injured or killed by the caustic, it ceases to grow, whilst the opposite side continues growing; and the result is that the tip itself bends towards the injured side and often becomes completely hooked; and it is remarkable that in this case the adjoining upper part does not bend. The stimulus is too powerful or the shock too great for the proper influence to be transmitted from the tip. We have strictly analogous cases with Drosera, Dionaea and Pinguicula, with which plants a too powerful stimulus does not excite the tentacles to become incurved, or the lobes to close, or the margin to be folded inwards.
With respect to the degree of sensitiveness of the apex to contact under favourable conditions, we have seen that with Vicia faba a little square of writing-paper affixed with shellac sufficed to cause movement; as did on one occasion a square of merely damped goldbeaters’ skin, but it acted very slowly. Short bits of moderately thick bristle (of which measurements have been given) affixed with gum-water acted in only three out of eleven trials, and beads of dried shellac under 1/200th of a grain in weight acted only twice in nine cases; so that here we have nearly reached the minimum of necessary irritation. The apex, therefore, is much less sensitive to pressure than the glands of Drosera, for these are affected by far thinner objects than bits of bristle, and by a very much less weight than 1/200th of a grain.
But the most interesting evidence of the delicate sensitiveness of the tip of the radicle, was afforded by its power of discriminating between equal-sized squares of card-like and very thin paper, when these were attached on opposite sides, as was observed with the radicles of the bean and oak.
When radicles of the bean are extended horizontally with squares of card attached to the lower sides of their tips, the irritation thus caused was always conquered by geotropism, which then acts under the most favourable conditions at right angles to the radicle. But when objects were attached to the radicles of any of the above-named genera, suspended vertically, the irritation conquered geotropism, which latter power at first acted obliquely on the radicle; so that the immediate irritation from the attached object, aided by its after-effects, prevailed and caused the radicle to bend upwards, until sometimes the point was directed to the zenith. We must, however, assume that the after-effects of the irritation of the tip by an attached object come into play, only after movement has been excited. The tips of the radicles of the pea seem to be more sensitive to contact than those of the bean, for when they were extended horizontally with squares of card adhering to their lower sides, a most curious struggle occasionally arose, sometimes one and sometimes the other force prevailing, but ultimately geotropism was always victorious; nevertheless, in two instances the terminal part became so much curved upwards that loops were subsequently formed. With the pea, therefore, the irritation from an attached object, and from geotropism when acting at right angles to the radicle, are nearly balanced forces. Closely similar results were observed with the horizontally extended radicles of _Cucurbita ovifera_, when their tips were slightly cauterised on the lower side.
Finally, the several co-ordinated movements by which radicles are enabled to perform their proper functions are admirably perfect. In whatever direction the primary radicle first protrudes from the seed, geotropism guides it perpendicularly downwards; and the capacity to be acted on by the attraction of gravity resides in the tip. But Sachs has proved[15] that the secondary radicles, or those emitted by the primary one, are acted on by geotropism in such a manner that they tend to bend only obliquely downwards. If they had been acted on like the primary radicle, all the radicles would have penetrated the ground in a close bundle. We have seen that if the end of the primary radicle is cut off or injured, the adjoining secondary radicles become geotropic and grow vertically downwards. This power must often be of great service to the plant, when the primary radicle has been destroyed by the larvae of insects, burrowing animals, or any other accident. The tertiary radicles, or those emitted by the secondary ones, are not influenced, at least in the case of the bean, by geotropism; so they grow out freely in all directions. From this manner of growth of the various kinds of radicles, they are distributed, together with their absorbent hairs, throughout the surrounding soil, as Sachs has remarked, in the most advantageous manner; for the whole soil is thus closely searched.
[15] ‘Arbeiten Bot. Institut, Würzburg,’ Heft iv. 1874, pp. 605–631.
Geotropism, as was shown in the last chapter, excites the primary radicle to bend downwards with very little force, quite insufficient to penetrate the ground. Such penetration is effected by the pointed apex (protected by the root-cap) being pressed down by the longitudinal expansion or growth of the terminal rigid portion, aided by its transverse expansion, both of which forces act powerfully. It is, however, indispensable that the seeds should be at first held down in some manner. When they lie on the bare surface they are held down by the attachment of the root-hairs to any adjoining objects; and this apparently is effected by the conversion of their outer surfaces into a cement. But many seeds get covered up by various accidents, or they fall into crevices or holes. With some seeds their own weight suffices.
The circumnutating movement of the terminal growing part both of the primary and secondary radicles is so feeble that it can aid them very little in penetrating the ground, excepting when the superficial layer is very soft and damp. But it must aid them materially when they happen to break obliquely into cracks, or into burrows made by earth-worms or larvae. This movement, moreover, combined with the sensitiveness of the tip to contact, can hardly fail to be of the highest importance; for as the tip is always endeavouring to bend to all sides it will press on all sides, and will thus be able to discriminate between the harder and softer adjoining surfaces, in the same manner as it discriminated between the attached squares of card-like and thin paper. Consequently it will tend to bend from the harder soil, and will thus follow the lines of least resistance. So it will be if it meets with a stone or the root of another plant in the soil, as must incessantly occur. If the tip were not sensitive, and if it did not excite the upper part of the root to bend away, whenever it encountered at right angles some obstacle in the ground, it would be liable to be doubled up into a contorted mass. But we have seen with radicles growing down inclined plates of glass, that as soon as the tip merely touched a slip of wood cemented across the plate, the whole terminal growing part curved away, so that the tip soon stood at right angles to its former direction; and thus it would be with an obstacle encountered in the ground, as far as the pressure of the surrounding soil would permit. We can also understand why thick and strong radicles, like those of Æsculus, should be endowed with less sensitiveness than more delicate ones; for the former would be able by the force of their growth to overcome any slight obstacle.
After a radicle, which has been deflected by some stone or root from its natural downward course, reaches the edge of the obstacle, geotropism will direct it to grow again straight downward; but we know that geotropism acts with very little force, and here another excellent adaptation, as Sachs has remarked,[16] comes into play. For the upper part of the radicle, a little above the apex, is, as we have seen, likewise sensitive; and this sensitiveness causes the radicle to bend like a tendril towards the touching object, so that as it rubs over the edge of an obstacle, it will bend downwards; and the curvature thus induced is abrupt, in which respect it differs from that caused by the irritation of one side of the tip. This downward bending coincides with that due to geotropism, and both will cause the root to resume its original course.
As radicles perceive an excess of moisture in the air on one side and bend towards this side, we may infer that they will act in the same manner with respect to moisture in the earth. The sensitiveness to moisture resides in the tip, which determines the bending of the upper part.
This capacity perhaps partly accounts for the extent to which drain-pipes often become choked with roots.
Considering the several facts given in this chapter, we see that the course followed by a root through the soil is governed by extraordinarily complex and diversified agencies,—by geotropism acting in a different manner on the primary, secondary, and tertiary radicles,—by sensitiveness to contact, different in kind in the apex and in the part immediately above the apex, and apparently by sensitiveness to the varying dampness of different parts of the soil.
These several stimuli to movement are all more powerful than geotropism, when this acts obliquely on a radicle, which has been deflected from its perpendicular downward course. The roots, moreover, of most plants are excited by light to bend either to or from it; but as roots are not naturally exposed to the light it is doubtful whether this sensitiveness, which is perhaps only the indirect result of the radicles being highly sensitive to other stimuli, is of any service to the plant. The direction which the apex takes at each successive period of the growth of a root, ultimately determines its whole course; it is therefore highly important that the apex should pursue from the first the most advantageous direction; and we can thus understand why sensitiveness to geotropism, to contact and to moisture, all reside in the tip, and why the tip determines the upper growing part to bend either from or to the exciting cause. A radicle may be compared with a burrowing animal such as a mole, which wishes to penetrate perpendicularly down into the ground. By continually moving his head from side to side, or circumnutating, he will feel any stone or other obstacle, as well as any difference in the hardness of the soil, and he will turn from that side; if the earth is damper on one than on the other side he will turn thitherward as a better hunting-ground. Nevertheless, after each interruption, guided by the sense of gravity, he will be able to recover his downward course and to burrow to a greater depth.
CHAPTER IV.
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.
We have seen in the first chapter that the stems of all seedlings, whether hypocotyls or epicotyls, as well as the cotyledons and the radicles, are continually circumnutating—that is they grow first on one side and then on another, such growth being probably preceded by increased turgescence of the cells. As it was unlikely that plants should change their manner of growth with advancing age, it seemed probable that the various organs of all plants at all ages, as long as they continued to grow, would be found to circumnutate, though perhaps to an extremely small extent. As it was important for us to discover whether this was the case, we determined to observe carefully a certain number of plants which were growing vigorously, and which were not known to move in any manner. We commenced with stems. Observations of this kind are tedious, and it appeared to us that it would be sufficient to observe the stems in about a score of genera, belonging to widely distinct families and inhabitants of various countries.
Several plants were selected which, from being woody, or for other reasons, seemed the least likely to circumnutate. The observations and the diagrams were made in the manner described in the Introduction. Plants in pots were subjected to a proper temperature, and whilst being observed, were kept either in darkness or were feebly illuminated from above. They are arranged in the order adopted by Hooker in Le Maout and Decaisne’s ‘System of Botany.’ The number of the family to which each genus belongs is appended, as this serves to show the place of each in the series.
(1.) Iberis umbellata (Cruciferae, Fam. 14).—The movement of the stem of a young plant, 4 inches in height, consisting of four internodes (the hypocotyl included) besides a large bud on the summit, was traced, as here shown, during 24 h. (Fig. 70). As far as we could judge the uppermost inch alone of the stem circumnutated, and this in a simple manner. The movement was slow, and the rate very unequal at different times. In part of its course an irregular ellipse, or rather triangle, Fig. 70. Iberis umbellata: circumnutation of stem of young plant, traced from 8.30 A.M. Sept. 13th to same hour on following morning.
Distance of summit of stem beneath the horizontal glass 7.6 inches.
Diagram reduced to half of original size. Movement as here shown magnified between 4 and 5 times.
(2.) Brassica oleracea (Cruciferae).—A very young plant, bearing three leaves, of which the longest was only three-quarters of an inch in length, was placed under a microscope, furnished with an eye-piece micrometer, and the tip of the largest leaf was found to be in constant movement. It crossed five divisions of the micrometer, that is, 1/100th of an inch, in 6 m. 20 s. There could hardly be a doubt that it was the stem which chiefly moved, for the tip did not get quickly out of focus; and this would have occurred had the movement been confined to the leaf, which moves up or down in nearly the same vertical plane.
(3.) Linum usitatissimum (Lineae, Fam. 39).—The stems of this plant, shortly before the flowering period, are stated by Fritz Müller (4.) Pelargonium zonale (Geraniaceae, Fam. 47).—A young plant, 7½ inches in height, was observed in the usual manner; but, in order to see the bead at the end of the glass filament and at the same time the mark beneath, it was necessary to cut off three leaves on one side. We do not know whether it was owing to this cause, or to the plant having previously become bent to one side through heliotropism, but from the morning of the 7th of March to 10.30 P.M. on the 8th, the stem moved a considerable distance in a zigzag line in the same general direction.
During the night of the 8th it moved to some distance at right angles to its former course, and next morning (9th) stood for a time almost still. At noon on the 9th a new tracing was begun (see Fig. 71), which was continued till 8 A.M. on the 11th. Between noon on the 9th and 5