SigPhi · Charles Darwin

The Power of Movement in Plants

Page 9 of 32

paragraphs) are here included. Out of 55 radicles experimented on under the proper temperature, 52 became bent, generally to a considerable extent from the perpendicular, and away from the side to which the object was attached. Of the three failures, one can be accounted for, as the radicle became sickly on the following day; and a second was observed only during 11 h. 30 m. As in several cases the terminal growing part of the radicle continued for some time to bend from the attached object, it formed itself into a hook, with the apex pointing to the zenith, or even into a ring, and occasionally into a spire or helix. It is remarkable that these latter cases occurred more frequently when objects were attached with thick gum-water, which never became dry, than when shellac was employed. The curvature was often well-marked in from 7 h. to 11 h.; and in one instance a semicircle was formed in 6 h. 10 m, from the time of attachment. But in order to see the phenomenon as well displayed as in the above described cases, it is indispensable that the bits of card, etc., should be made to adhere closely to one side of the conical apex; that healthy radicles should be selected and kept at not too high or too low a temperature, and apparently that the trials should not be made in the middle of the winter.

In ten instances, radicles which had curved away from a square of card or other object attached to their tips, straightened themselves to a certain extent, or even completely, in the course of from one to two days from the time of attachment. This was more especially apt to occur when the curvature was slight. But in one instance (No. 27) a radicle which in 9 h. had been deflected about 90° from the perpendicular, became quite straight in 24 h. from the period of attachment. With No.

26, the radicle was almost straight in 48 h. We at first attributed the straightening process to the radicles becoming accustomed to a slight stimulus, in the same manner as a tendril or sensitive petiole becomes accustomed to a very light loop of thread, and unbends itself though the loop remains still suspended; but Sachs states[2] that radicles of the bean placed horizontally in damp air after curving downwards through geotropism, straighten themselves a little by growth along their lower or concave sides. Why this should occur is not clear: but perhaps it likewise occurred in the above ten cases. There is another occasional movement which must not be passed over: the tip of the radicle, for a length of from 2 to 3 mm., was found in six instances, after an interval of about 24 or more hours, bent towards the bit of still attached card,—that is, in a direction exactly opposite to the previously induced curvature of the whole growing part for a length of from 7 to 8 mm. This occurred chiefly when the first curvature was small, and when an object had been affixed more than once to the apex of the same radicle. The attachment of a bit of card by shellac to one side of the tender apex may sometimes mechanically prevent its growth; or the application of thick gum-water more than once to the same side may injure it; and then checked growth on this side with continued growth on the opposite and unaffected side would account for the reversed curvature of the apex.

Various trials were made for ascertaining, as far as we could, the nature and degree of irritation to which the apex must be subjected, in order that the terminal growing part should bend away, as if to avoid the cause of irritation. We have seen in the numbered experiments, that a little square of rather thick letter-paper gummed to the apex induced, though slowly, considerable deflection. Judging from several cases in which various objects had been affixed with gum, and had soon become separated from the apex by a layer of fluid, as well as from some trials in which drops of thick gum-water alone had been applied, this fluid never causes bending. We have also seen in the numbered experiments that narrow splinters of quill and of very thin glass, affixed with shellac, caused only a slight degree of deflection, and this may perhaps have been due to the shellac itself. Little squares of goldbeaters’ skin, which is excessively thin, were damped, and thus made to adhere to one side of the tips of two radicles; one of these, after 24 h., produced no effect; nor did the other in 8 h., within which time squares of card usually act; but after 24 h. there was slight deflection.

An oval bead, or rather cake, of dried shellac, 1.01 mm. in length and 0.63 in breadth, caused a radicle to become deflected at nearly right angles in the course of only 6 h.; but after 23 h. it had nearly straightened itself. A very small quantity of dissolved shellac was spread over a bit of card, and the tips of 9 radicles were touched laterally with it; only two of them became slightly deflected to the side opposite to that bearing the speck of dried shellac, and they afterwards straightened themselves. These specks were removed, and both together weighed less than 1/100th of a grain; so that a weight of rather less than 1/200th of a grain (0.32 mg.) sufficed to excite movement in two out of the nine radicles. Here then we have apparently reached nearly the minimum weight which will act.

A moderately thick bristle (which on measurement was found rather flattened, being 0.33 mm. in one diameter, and 0.20 mm. in the other) was cut into lengths of about 1/20th of an inch. These after being touched with thick gum-water, were placed on the tips of eleven radicles. Three of them were affected; one being deflected in 8 h. 15 m. to an angle of about 90° from the perpendicular; a second to the same amount when looked at after 9 h.; but after 24 h. from the time of first attachment the deflection had decreased to only 19°; the third was only slightly deflected after 9 h., and the bit of bristle was then found not touching the apex; it was replaced, and after 15 additional hours the deflection amounted to 26° from the perpendicular. The remaining eight radicles were not at all acted on by the bits of bristle, so that we here appear to have nearly reached the minimum of size of an object which will act on the radicle of the bean. But it is remarkable that when the bits of bristle did act, that they should have acted so quickly and efficiently.

As the apex of a radicle in penetrating the ground must be pressed on all sides, we wished to learn whether it could distinguish between harder or more resisting, and softer substances. A square of the sanded paper, almost as stiff as card, and a square of extremely thin paper (too thin for writing on), of exactly the same size (about 1/20th of an inch), were fixed with shellac on opposite sides of the apices of 12 suspended radicles. The sanded card was between 0.15 and 0.20 mm. (or between 0.0059 and 0.0079 of an inch), and the thin paper only 0.045 mm. (or 0.00176 of an inch) in thickness. In 8 out of the 12 cases there could be no doubt that the radicle was deflected from the side to which the card-like paper was attached, and towards the opposite side, bearing the very thin paper. This occurred in some instances in 9 h., but in others not until 24 h. had elapsed. Moreover, some of the four failures can hardly be considered as really failures: thus, in one of them, in which the radicle remained quite straight, the square of thin paper was found, when both were removed from the apex, to have been so thickly coated with shellac that it was almost as stiff as the card: in the second case, the radicle was bent upwards into a semicircle, but the deflection was not directly from the side bearing the card, and this was explained by the two squares having become cemented laterally together, forming a sort of stiff gable, from which the radicle was deflected: in the third case, the square of card had been fixed by mistake in front, and though there was deflection from it, this might have been due to Sachs’ curvature: in the fourth case alone no reason could be assigned why the radicle had not been at all deflected. These experiments suffice to prove that the apex of the radicle possesses the extraordinary power of discriminating between thin card and very thin paper, and is deflected from the side pressed by the more resisting or harder substance.

Some trials were next made by irritating the tips without any object being left in contact with them. Nine radicles, suspended over water, had their tips rubbed, each six times with a needle, with sufficient force to shake the whole bean; the temperature was favourable, viz.

about 63° F. In 7 out of these cases no effect whatever was produced; in the eighth case the radicle became slightly deflected from, and in the ninth case slightly deflected towards, the rubbed side; but these two latter opposed curvatures were probably accidental, as radicles do not always grow perfectly straight downwards. The tips of two other radicles were rubbed in the same manner for 15 seconds with a little round twig, two others for 30 seconds, and two others for 1 minute, but without any effect being produced. We may therefore conclude from these 15 trials that the radicles are not sensitive to temporary contact, but are acted on only by prolonged, though very slight, pressure.

We then tried the effects of cutting off a very thin slice parallel to one of the sloping sides of the apex, as we thought that the wound would cause prolonged irritation, which might induce bending towards the opposite side, as in the case of an attached object. Two preliminary trials were made: firstly, slices were cut from the radicles of 6 beans suspended in damp air, with a pair of scissors, which, though sharp, probably caused considerable crushing, and no curvature followed. Secondly, thin slices were cut with a razor obliquely off the tips of three radicles similarly suspended; and after 44 h. two were found plainly bent from the sliced surface; and the third, the whole apex of which had been cut off obliquely by accident, was curled upwards over the bean, but it was not clearly ascertained whether the curvature had been at first directed from the cut surface.

These results led us to pursue the experiment, and 18 radicles, which had grown vertically downwards in damp air, had one side of their conical tips sliced off with a razor. The tips were allowed just to enter the water in the jars, and they were exposed to a temperature 14°–16° C. (57°–61° F.). The observations were made at different times.

Three were examined 12 h. after being sliced, and were all slightly curved from the cut surface; and the curvature increased considerably after an additional 12 h. Eight were examined after 19 h.; four after 22 h. 30 m.; and three after 25 h. The final result was that out of the 18 radicles thus tried, 13 were plainly bent from the cut surface after the above intervals of time; and one other became so after an additional interval of 13 h. 30 m. So that only 4 out of the 18 radicles were not acted on. To these 18 cases the 3 previously mentioned ones should be added. It may, therefore, be concluded that a thin slice removed by a razor from one side of the conical apex of the radicle causes irritation, like that from an attached object, and induces curvature from the injured surface.

Lastly, dry caustic (nitrate of silver) was employed to irritate one side of the apex. If one side of the apex or of the whole terminal growing part of a radicle, is by any means killed or badly injured, the other side continues to grow; and this causes the part to bend over towards the injured side.[3] But in the following experiments we endeavoured, generally with success, to irritate the tips on one side, without badly injuring them. This was effected by first drying the tip as far as possible with blotting-paper, though it still remained somewhat damp, and then touching it once with quite dry caustic. Seventeen radicles were thus treated, and were suspended in moist air over water at a temperature of 58° F. They were examined after an interval of 21 h. or 24h. The tips of two were found blackened equally all round, so that they could tell nothing and were rejected, 15 being left. Of these, 10 were curved from the side which had been touched, where there was a minute brown or blackish mark. Five of these radicles, three of which were already slightly deflected, were allowed to enter the water in the jar, and were re-examined after an additional interval of 27 h. (i.e. in 48 h. after the application of the caustic), and now four of them had become hooked, being bent from the discoloured side, with their points directed to the zenith; the fifth remained unaffected and straight. Thus 11 radicles out of the 15 were acted on.

But the curvature of the four just described was so plain, that they alone would have sufficed to show that the radicles of the bean bend away from that side of the apex which has been slightly irritated by caustic.

[3] Ciesielski found this to be the case (‘Untersuchungen über die Abwartskrümmung der Wurzel,’ 1871, p. 28) after burning with heated platinum one side of a radicle. So did we when we painted longitudinally half of the whole length of 7 radicles, suspended over water, with a thick layer of grease, which is very injurious or even fatal to growing parts; for after 48 hours five of these radicles were curved towards the greased side, two remaining straight.

_The Power of an Irritant on the apex of the Radicle_ _of the Bean, compared with that of Geotropism_.—We know that when a little square of card or other object is fixed to one side of the tip of a vertically dependent radicle, the growing part bends from it often into a semicircle, in opposition to geotropism, which force is conquered by the effect of the irritation from the attached object.

Radicles were therefore extended horizontally in damp air, kept at the proper low temperature for full sensitiveness, and squares of card were affixed with shellac on the lower sides of their tips, so that if the squares acted, the terminal growing part would curve upwards. Firstly, eight beans were so placed that their short, young, horizontally extended radicles would be simultaneously acted on both by geotropism and by Sachs’ curvature, if the latter came into play; and they all eight became bowed downwards to the centre of the earth in 20 h., excepting one which was only slightly acted on. Two of them were a little bowed downwards in only 5 h.! Therefore the cards, affixed to the lower sides of their tips, seemed to produce no effect; and geotropism easily conquered the effects of the irritation thus caused.

Secondly, 5 oldish radicles, 1½ inch in length, and therefore less sensitive than the above-mentioned young ones, were similarly placed and similarly treated. From what has been seen on many other occasions, it may be safely inferred that if they had been suspended vertically they would have bent away from the cards; and if they had been extended horizontally, without cards attached to them, they would have quickly bent vertically downwards through geotropism; but the result was that two of these radicles were still horizontal after 23 h.; two were curved only slightly, and the fifth as much as 40° beneath the horizon.

Thirdly, 5 beans were fastened with their flat surfaces parallel to the cork-lid, so that Sachs’ curvature would not tend to make the horizontally extended radicles turn either upwards or downwards, and little squares of card were affixed as before, to the lower sides of their tips. The result was that all five radicles were bent down, or towards the centre of the earth, after only 8 h. 20 m. At the same time and within the same jars, 3 radicles of the same age, with squares affixed to one side, were suspended vertically; and after 8 h. 20 m. they were considerably deflected from the cards, and therefore curved upwards in opposition to geotropism. In these latter cases the irritation from the squares had over-powered geotropism; whilst in the former cases, in which the radicles were extended horizontally, geotropism had overpowered the irritation. Thus within the same jars, some of the radicles were curving upwards and others downwards at the same time—these opposite movements depending on whether the radicles, when the squares were first attached to them, projected vertically down, or were extended horizontally. This difference in their behaviour seems at first inexplicable, but can, we believe, be simply explained by the difference between the initial power of the two forces under the above circumstances, combined with the well-known principle of the after-effects of a stimulus. When a young and sensitive radicle is extended horizontally, with a square attached to the lower side of the tip, geotropism acts on it at right angles, and, as we have seen, is then evidently more efficient than the irritation from the square; and the power of geotropism will be strengthened at each successive period by its previous action—that is, by its after-effects. On the other hand, when a square is affixed to a vertically dependent radicle, and the apex begins to curve upwards, this movement will be opposed by geotropism acting only at a very oblique angle, and the irritation from the card will be strengthened by its previous action. We may therefore conclude that the initial power of an irritant on the apex of the radicle of the bean, is less than that of geotropism when acting at right angles, but greater than that of geotropism when acting obliquely on it.

Sensitiveness of the tips of the Secondary Radicles of the Bean to contact.—All the previous observations relate to the main or primary radicle. Some beans suspended to cork-lids, with their radicles dipping into water, had developed secondary or lateral radicles, which were afterwards kept in very damp air, at the proper low temperature for full sensitiveness. They projected, as usual, almost horizontally, with only a slight downward curvature, and retained this position during several days. Sachs has shown[4] that these secondary roots are acted on in a peculiar manner by geotropism, so that if displaced they reassume their former sub-horizontal position, and do not bend vertically downwards like the primary radicle. Minute squares of the stiff sanded paper were affixed by means of shellac (but in some instances with thick gum-water) to the tips of 39 secondary radicles of different ages, generally the uppermost ones. Most of the squares were fixed to the lower sides of the apex, so that if they acted the radicle would bend upwards; but some were fixed laterally, and a few on the upper side. Owing to the extreme tenuity of these radicles, it was very difficult to attach the square to the actual apex. Whether owing to this or some other circumstance, only nine of the squares induced any curvature. The curvature amounted in some cases to about 45° above the horizon, in others to 90°, and then the tip pointed to the zenith. In one instance a distinct upward curvature was observed in 8 h. 15 m., but usually not until 24 h. had elapsed. Although only 9 out of 39 radicles were affected, yet the curvature was so distinct in several of them, that there could be no doubt that the tip is sensitive to slight contact, and that the growing part bends away from the touching object.

It is possible that some secondary radicles are more sensitive than others; for Sachs has proved[5] the interesting fact that each individual secondary radicle possesses its own peculiar constitution.

Sensitiveness to contact of the Primary Radicle, a little above the apex, in the Bean (Vicia faba) and Pea (Pisum sativum).—The sensitiveness of the apex of the radicle in the previously described cases, and the consequent curvature of the upper part from the touching object or other source of irritation, is the more remarkable, because Sachs[6] has shown that pressure at the distance of a few millimeters above the apex causes the radicle to bend, like a tendril, towards the touching object. By fixing pins so that they pressed against the radicles of beans suspended vertically in damp air, we saw this kind of curvature; but rubbing the part with a twig or needle for a few minutes produced no effect. Haberlandt remarks,[7] that these radicles in breaking through the seed-coats often rub and press against the ruptured edges, and consequently bend round them. As little squares of the card-like paper affixed with shellac to the tips were highly efficient in causing the radicles to bend away from them, similar pieces (of about 1/20th inch square, or rather less) were attached in the same manner to one side of the radicle at a distance of 3 or 4 mm. above the apex. In our first trial on 15 radicles no effect was produced. In a second trial on the same number, three became abruptly curved (but only one strongly) towards the card within 24 h. From these cases we may infer that the pressure from a bit of card affixed with shellac to one side above the apex, is hardly a sufficient irritant; but that it occasionally causes the radicle to bend like a tendril towards this side.

We next tried the effect of rubbing several radicles at a distance of 4 mm. from the apex for a few seconds with lunar caustic (nitrate of silver); and although the radicles had been wiped dry and the stick of caustic was dry, yet the part rubbed was much injured and a slight permanent depression was left. In such cases the opposite side continues to grow, and the radicle necessarily becomes bent towards the injured side. But when a point 4 mm. from the apex was momentarily touched with dry caustic, it was only faintly discoloured, and no permanent injury was caused. This was shown by several radicles thus treated straightening themselves after one or two days; yet at first they became curved towards the touched side, as if they had been there subjected to slight continued pressure. These cases deserve notice, because when one side of the apex was just touched with caustic, the radicle, as we have seen, curved itself in an opposite direction, that is, away from the touched side.

The radicle of the common pea at a point a little above the apex is rather more sensitive to continued pressure than that of the bean, and bends towards the pressed side.[8] We experimented on a variety (_Yorkshire Hero_) which has a much wrinkled tough skin, too large for the included cotyledons; so that out of 30 peas which had been soaked for 24 h. and allowed to germinate on damp sand, the radicles of three were unable to escape, and were crumpled up in a strange manner within the skin; four other radicles were abruptly bent round the edges of the ruptured skin against which they had pressed. Such abnormalities would probably never, or very rarely, occur with forms developed in a state of nature and subjected to natural selection. One of the four radicles just mentioned in doubling backwards came into contact with the pin by which the pea was fixed to the cork-lid; and now it bent at right angles round the pin, in a direction quite different from that of the first curvature due to contact with the ruptured skin; and it thus afforded a good illustration of the tendril-like sensitiveness of the radicle a little above the apex.

[8] Sachs, ‘Arbeiten Bot. Institut., Würzburg,’ Heft iii. p. 438.

Little squares of the card-like paper were next affixed to radicles of the pea at 4 mm. above the apex, in the same manner as with the bean.

Twenty-eight radicles suspended vertically over water were thus treated on different occasions, and 13 of them became curved towards the cards.

The greatest degree of curvature amounted to 62° from the perpendicular; but so large an angle was only once formed. On one occasion a slight curvature was perceptible after 5 h. 45 m., and it was generally well-marked after 14 h. There can therefore be no doubt that with the pea, irritation from a bit of card attached to one side of the radicle above the apex suffices to induce curvature.

Squares of card were attached to one side of the tips of 11 radicles within the same jars in which the above trials were made, and five of them became plainly, and one slightly, curved away from this side.

Other analogous cases will be immediately described. The fact is here mentioned because it was a striking spectacle, showing the difference in the sensitiveness of the radicle in different parts, to behold in the same jar one set of radicles curved away from the squares on their tips, and another set curved towards the squares attached a little higher up. Moreover, the kind of curvature in the two cases is different. The squares attached above the apex cause the radicle to bend abruptly, the part above and beneath remaining nearly straight; so that here there is little or no transmitted effect. On the other hand, the squares attached to the apex affect the radicle for a length of from about 4 to even 8 mm., inducing in most cases a symmetrical curvature; so that here some influence is transmitted from the apex for this distance along the radicle.

Pisum sativum (var. Yorkshire Hero): Sensitiveness of the apex of the Radicle.—Little squares of the same card-like paper were affixed (April 24th) with shellac to one side of the apex of 10 vertically suspended radicles: the temperature of the water in the bottom of the jars was 60°–61° F. Most of these radicles were acted on in 8 h. 30 m.; and eight of them became in the course of 24 h. conspicuously, and the remaining two slightly, deflected from the perpendicular and from the side bearing the attached squares. Thus all were acted on; but it will suffice to describe two conspicuous cases. In one the terminal portion of the radicle was bent at right angles (A, Fig. 66) after 24h.; and in the other (B) it had by this time become hooked, with the apex pointing to the zenith. The two bits of card here used were.07 inch in length and.04 inch in breadth. Two other radicles, which after 8 h. 30 m.

were moderately deflected, became straight again after 24h. Another trial was made in the same manner with 15 radicles; but from circumstances, not worth explaining, they were only once and briefly examined after the short interval of 5 h. 30 m.; and we merely record in our notes “almost all bent slightly from the perpendicular, and away from the squares; the deflection amounting in one or two instances to nearly a rectangle.” These two sets of cases, especially the first one, prove that the apex of the radicle is sensitive to slight contact and that the upper part bends from the touching object. Nevertheless, on June 1st and 4th, 8 other radicles were tried in the same manner at a temperature of 58°–60° F., and after 24 h. only 1 was decidedly bent from the card, 4 slightly, 2 doubtfully, and 1 not in the least. The amount of curvature was unaccountably small; but all the radicles which were at all bent, were bent away from the cards.

Fig. 66. Pisum sativum: deflection produced within 24 hours in the growth of vertically dependent radicles, by little squares of card affixed with shellac to one side of apex: A, bent at right angles; B, hooked.

We now tried the effects of widely different temperatures on the sensitiveness of these radicles with squares of card attached to their tips. Firstly, 13 peas, most of them having very short and young radicles, were placed in an ice-box, in which the temperature rose during three days from 44° to 47° F. They grew slowly, but 10 out of the 13 became in the course of the three days very slightly curved from the squares; the other 3 were not affected; so that this temperature was too low for any high degree of sensitiveness or for much movement. Jars with 13 other radicles were next placed on a chimney-piece, where they were subjected to a temperature of between 68° and 72° F., and after 24 h., 4 were conspicuously curved from the cards, 2 slightly, and 7 not at all; so that this temperature was rather too high. Lastly 12 radicles were subjected to a temperature varying between 72° and 85° F., and none of them were in the least affected by the squares. The above several trials, especially the first recorded one, indicate that the most favourable temperature for the sensitiveness of the radicle of the pea is about 60° F.

The tips of 6 vertically dependent radicles were touched once with dry caustic, in the manner described under Vicia faba. After 24 h. four of them were bent from the side bearing a minute black mark; and the curvature increased in one case after 38 h., and in another case after 48 h., until the terminal part projected almost horizontally. The two remaining radicles were not affected.

With radicles of the bean, when extended horizontally in damp air, geotropism always conquered the effects of the irritation caused by squares of card attached to the lower sides of their tips. A similar experiment was tried on 13 radicles of the pea; the squares being attached with shellac, and the temperature between 58°–60° F. The result was somewhat different; for these radicles are either less strongly acted on by geotropism, or, what is more probable, are more sensitive to contact. After a time geotropism always prevailed, but its action was often delayed; and in three instances there was a most curious struggle between geotropism and the irritation caused by the cards. Four of the 13 radicles were a little curved downwards within 6 or 8 h., always reckoning from the time when the squares were first attached, and after 23 h. three of them pointed vertically downwards, and the fourth at an angle of 45° beneath the horizon. These four radicles therefore did not seem to have been at all affected by the attached squares. Four others were not acted on by geotropism within the first 6 or 8 h., but after 23 h. were much bowed down. Two others remained almost horizontal for 23 h., but afterwards were acted on. So that in these latter six cases the action of geotropism was much delayed. The eleventh radicle was slightly curved down after 8 h., but when looked at again after 23 h. the terminal portion was curved upwards; if it had been longer observed, the tip no doubt would have been found again curved down, and it would have formed a loop as in the following case.

The twelfth radicle after 6 h. was slightly curved downwards; but when looked at again after 21 h., this curvature had disappeared and the apex pointed upwards; after 30 h. the radicle formed a hook, as shown at A (Fig. 67); which hook after 45 h. was converted into a loop (B).

The thirteenth radicle after 6 h. was slightly curved downwards, but within 21 h. had curved considerably up, and then down again at an angle of 45° beneath the horizon, afterwards becoming perpendicular. In these three last cases geotropism and the irritation caused by the attached squares alternately prevailed in a highly remarkable manner; geotropism being ultimately victorious.

Fig. 67. Pisum sativum: a radicle extended horizontally in damp air with a little square of card affixed to the lower side of its tip, causing it to bend upwards in opposition to geotropism. The deflection of the radicle after 21 hours is shown at A, and of the same radicle after 45 hours at B, now forming a loop.

Similar experiments were not always quite so successful as in the above cases. Thus 6 radicles, horizontally extended with attached squares, were tried on June 8th at a proper temperature, and after 7 h. 30 m.

none were in the least curved upwards and none were distinctly geotropic; whereas of 6 radicles without any attached squares, which served as standards of comparison or controls, 3 became slightly and 3 almost rectangularly geotropic within the 7 h. 30 m.; but after 23 h.

the two lots were equally geotropic. On July 10th another trial was made with 6 horizontally extended radicles, with squares attached in the same manner beneath their tips; and after 7 h. 30 m., 4 were slightly geotropic, 1 remained horizontal, and 1 was curved upwards in opposition to gravity or geotropism. This latter radicle after 48 h.

formed a loop, like that at B (Fig. 67).

An analogous trial was now made, but instead of attaching squares of card to the lower sides of the tips, these were touched with dry caustic. The details of the experiment will be given in the chapter on Geotropism, and it will suffice here to say that 10 peas, with radicles extended horizontally and not cauterised, were laid on and under damp friable peat; these, which served as standards or controls, as well as 10 others which had been touched on the upper side with the caustic, all became strongly geotropic in 24 h. Nine radicles, similarly placed, had their tips touched on the lower side with the caustic; and after 24 h., 3 were slightly geotropic, 2 remained horizontal, and 4 were bowed upwards in opposition to gravity and to geotropism. This upward curvature was distinctly visible in 8 h. 45m. after the lower sides of the tips had been cauterised.

Little squares of card were affixed with shellac on two occasions to the tips of 22 young and short secondary radicles, which had been emitted from the primary radicle whilst growing in water, but were now suspended in damp air. Besides the difficulty of attaching the squares to such finely pointed objects as were these radicles, the temperature was too high,—varying on the first occasion from 72° to 77° F., and on the second being almost steadily 78° F.; and this probably lessened the sensitiveness of the tips. The result was that after an interval of 8 h. 30 m., 6 of the 22 radicles were bowed upwards (one of them greatly) in opposition to gravity, and 2 laterally; the remaining 14 were not affected. Considering the unfavourable circumstances, and bearing in mind the case of the bean, the evidence appears sufficient to show that the tips of the secondary radicles of the pea are sensitive to slight contact.

Phaseolus multiflorus: Sensitiveness of the apex of the Radicle.—Fifty-nine radicles were tried with squares of various sizes of the same card-like paper, also with bits of thin glass and rough cinders, affixed with shellac to one side of the apex.

Rather large drops of the dissolved shellac were also placed on them and allowed to set into hard beads. The specimens were subjected to various temperatures between 60° and 72° F., more commonly at about the latter. But out of this considerable number of trials only 5 radicles were plainly bent, and 8 others slightly or even doubtfully, from the attached objects; the remaining 46 not being at all affected. It is therefore clear that the tips of the radicles of this Phaseolus are much less sensitive to contact than are those of the bean or pea. We thought that they might be sensitive to harder pressure, but after several trials we could not devise any method for pressing harder on one side of the apex than on the other, without at the same time