We were much more successful with caustic; but in our first trial, 15 radicles were too much cauterised, and only two became curved from the blackened side; the others being either killed on one side, or blackened equally all round. In our next trial the dried tips of 11 radicles were touched momentarily with dry caustic, and after a few minutes were immersed in water. The elongated marks thus caused were never black, only brown, and about mm. in length, or even less. In 4 h. 30 m. after the cauterisation, 6 of them were plainly curved from the side with the brown mark, 4 slightly, and 1 not at all. The latter proved unhealthy, and never grew; and the marks on 2 of the 4 slightly curved radicles were excessively minute, one being distinguishable only with the aid of a lens. Of 10 control specimens tried in the same jars at the same time, not one was in the least curved. In 8 h. 40 m. after the cauterisation, 5 of the radicles out of the 10 (the one unhealthy one being omitted) were deflected at about 90o, and 3 at about 45o from the perpendicular and from the side bearing the brown mark. After 24 h. all 10 radicles had increased immensely in length; in 5 of them the curvature was nearly the same, in 2 it had increased, and in 3 it had decreased. The contrast presented by the 10 controls, after both the 8 h. 40 m. and the 24 h. intervals, was very great; for they had continued to grow vertically downwards, excepting two which, from some unknown cause, had become somewhat tortuous.
In the chapter on Geotropism we shall see that 10 radicles of this plant were extended horizontally on and beneath damp friable peat, under which conditions [page 171]
they grow better and more naturally than in damp air; and their tips were slightly cauterised on the lower side, brown marks about mm. in length being thus caused. Uncauterised specimens similarly placed became much bent downwards through geotropism in the course of 5 or 6 hours. After 8 h. only 3 of the cauterised ones were bowed downwards, and this in a slight degree; 4 remained horizontal; and 3 were curved upwards in opposition to geotropism and from the side bearing the brown mark. Ten other specimens had their tips cauterised at the same time and in the same degree, on the upper side; and this, if it produced any effect, would tend to increase the power of geotropism; and all these radicles were strongly bowed downwards after 8 h. From the several foregoing facts, there can be no doubt that the cauterisation of the tip of the radicle of this Cucurbita on one side, if done lightly enough, causes the whole growing part to bend to the opposite side.
Rapha.n.u.s sativus: Sensitiveness of the apex of the Radicle.--We here encountered many difficulties in our trials, both with squares of card and with caustic; for when seeds were pinned to a cork-lid, many of the radicles, to which nothing had been done, grew irregularly, often curving upwards, as if attracted by the damp surface above; and when they were immersed in water they likewise often grew irregularly. We did not therefore dare to trust our experiments with attached squares of card; nevertheless some of them seemed to indicate that the tips were sensitive to contact. Our trials with caustic generally failed from the difficulty of not injuring too greatly the extremely fine tips. Out of 7 radicles thus tried, one became bowed after 22 h. at an angle of 60o, a second at 40o, [page 172]
and a third very slightly from the perpendicular and from the cauterised side.
Aesculus hippocastanum: Sensitiveness of the apex of the Radicle.--Bits of gla.s.s and squares of card were affixed with sh.e.l.lac or gum-water to the tips of 12 radicles of the horse-chestnut; and when these objects fell off, they were refixed; but not in a single instance was any curvature thus caused. These ma.s.sive radicles, one of which was above 2 inches in length and .3 inch in diameter at its base, seemed insensible to so slight a stimulus as any small attached object. Nevertheless, when the apex encountered an obstacle in its downward course, the growing part became so uniformly and symmetrically curved, that its appearance indicated not mere mechanical bending, but increased growth along the whole convex side, due to the irritation of the apex.
That this is the correct view may be inferred from the effects of the more powerful stimulus of caustic. The bending from the cauterised side occurred much slower than in the previously described species, and it will perhaps be worth while to give our trials in detail.
[The seeds germinated in sawdust, and one side of the tips of the radicles were slightly rubbed once with dry nitrate of silver; and after a few minutes were allowed to dip into water. They were subjected to a rather varying temperature, generally between 52o and 58o F. A few cases have not been thought worth recording, in which the whole tip was blackened, or in which the seedling soon became unhealthy.
(1.) The radicle was slightly deflected from the cauterised side in one day (i.e. 24 h.); in three days it stood at 60o from the perpendicular; in four days at 90o; on the fifth day it was curved up about 40o above the horizon; so that it had pa.s.sed through an angle of 130o in the five days, and this was the greatest amount of curvature observed.
(2.) In two days radicle slightly deflected; after seven days [page 173]
deflected 69o from the perpendicular and from the cauterised side; after eight days the angle amounted to nearly 90o.
(3.) After one day slight deflection, but the cauterised mark was so faint that the same side was again touched with caustic. In four days from the first touch deflection amounted to 78o, which in an additional day increased to 90o.
(4.) After two days slight deflection, which during the next three days certainly increased but never became great; the radicle did not grow well and died on the eighth day.
(5.) After two days very slight deflection; but this on the fourth day amounted to 56o from the perpendicular and from the cauterised side.
(6.) After three days doubtfully, but after four days certainly deflected from the cauterised side. On the fifth day deflection amounted to 45o from the perpendicular, and this on the seventh day increased to about 90o.
(7.) After two days slightly deflected; on the third day the deflection amounted to 25o from the perpendicular, and this did not afterwards increase.
(8.) After one day deflection distinct; on the third day it amounted to 44o, and on the fourth day to 72o from the perpendicular and the cauterised side.
(9.) After two days deflection slight, yet distinct; on the third day the tip was again touched on the same side with caustic and thus killed.
(10.) After one day slight deflection, which after six days increased to 50o from the perpendicular and the cauterised side.
(11.) After one day decided deflection, which after six days increased to 62o from the perpendicular and from the cauterised side.
(12.) After one day slight deflection, which on the second day amounted to 35o, on the fourth day to 50o, and the sixth day to 63o from the perpendicular and the cauterised side.
(13.) Whole tip blackened, but more on one side than the other; on the fourth day slightly, and on the sixth day greatly deflected from the more blackened side; the deflection on the ninth day amounted to 90o from the perpendicular.
(14.) Whole tip blackened in the same manner as in the last case: on the second day decided deflection from the more blackened side, which increased on the seventh day to nearly 90o; on the following day the radicle appeared unhealthy.
(15.) Here we had the anomalous case of a radicle bending [page 174]
slightly towards the cauterised side on the first day, and continuing to do so for the next three days, when the deflection amounted to about 90o from the perpendicular. The cause appeared to lie in the tendril-like sensitiveness of the upper part of the radicle, against which the point of a large triangular flap of the seed-coats pressed with considerable force; and this irritation apparently conquered that from the cauterised apex.]
These several cases show beyond doubt that the irritation of one side of the apex, excites the upper part of the radicle to bend slowly towards the opposite side. This fact was well exhibited in one lot of five seeds pinned to the cork-lid of a jar; for when after 6 days the lid was turned upside down and viewed from directly above, the little black marks made by the caustic were now all distinctly visible on the upper sides of the tips of the laterally bowed radicles. A thin slice was shaved off with a razor from one side of the tips of 22 radicles, in the manner described under the common bean; but this kind of irritation did not prove very effective. Only 7 out of the 22 radicles became moderately deflected in from 3 to 5 days from the sliced surface, and several of the others grew irregularly. The evidence, therefore, is far from conclusive.
Quercus robur: Sensitiveness of the apex of the Radicle.--The tips of the radicles of the common oak are fully as sensitive to slight contact as are those of any plant examined by us. They remained healthy in damp air for 10 days, but grew slowly. Squares of the card-like paper were fixed with sh.e.l.lac to the tips of 15 radicles, and ten of these became conspicuously bowed from the perpendicular and from the squares; two slightly, and three not at all. But two of the latter were not real exceptions, as they were at first very short, and hardly grew afterwards. Some of the more [page 175]
remarkable cases are worth describing. The radicles were examined on each successive morning, at nearly the same hour, that is, after intervals of 24 h.
[No. 1. This radicle suffered from a series of accidents, and acted in an anomalous manner, for the apex appeared at first insensible and afterwards sensitive to contact. The first square was attached on Oct 19th; on the 21st the radicle was not at all curved, and the square was accidentally knocked off; it was refixed on the 22nd, and the radicle became slightly curved from the square, but the curvature disappeared on the 23rd, when the square was removed and refixed. No curvature ensued, and the square was again accidentally knocked off, and refixed. On the morning of the 27th it was washed off by having reached the water in the bottom of the jar. The square was refixed, and on the 29th, that is, ten days after the first square had been attached, and two days after the attachment of the last square, the radicle had grown to the great length of 3.2 inches, and now the terminal growing part had become bent away from the square into a hook (see Fig. 68).
Fig. 68. Quercus robur: radicle with square of card attached to one side of apex, causing it to become hooked. Drawing one-half natural scale.
No. 2. Square attached on the 19th; on the 20th radicle slightly deflected from it and from the perpendicular; on the 21st deflected at nearly right angles; it remained during the next two days in this position, but on the 25th the upward curvature was lessened through the action of geotropism, and still more so on the 26th.
No. 3. Square attached on the 19th; on the 21st a trace of curvature from the square, which amounted on the 22nd to about 40o, and on the 23rd to 53o from the perpendicular.
No. 4. Square attached on the 21st; on the 22nd trace of curvature from the square; on the 23rd completely hooked with the point turned up to the zenith. Three days afterwards (i.e. 26th) the curvature had wholly disappeared and the apex pointed perpendicularly downwards.
No. 5. Square attached on the 21st; on the 22nd decided [page 176]
though slight curvature from the square; on the 23rd the tip had curved up above the horizon, and on the 24th was hooked with the apex pointing almost to the zenith, as in Fig. 68.
No. 6. Square attached on the 21st; on the 22nd slightly curved from the square; 23rd more curved; 25th considerably curved; 27th all curvature lost, and the radicle was now directed perpendicularly downwards.
No. 7. Square attached on the 21st; on the 22nd a trace of curvature from the square, which increased next day, and on the 24th amounted to a right angle.
It is, therefore, manifest that the apex of the radicle of the oak is highly sensitive to contact, and retains its sensitiveness during several days. The movement thus induced was, however, slower than in any of the previous cases, with the exception of that of Aesculus. As with the bean, the terminal growing part, after bending, sometimes straightened itself through the action of geotropism, although the object still remained attached to the tip.
The same remarkable experiment was next tried, as in the case of the bean; namely, little squares of exactly the same size of the card-like sanded paper and of very thin paper (the thicknesses of which have been given under Vicia faba) were attached with sh.e.l.lac on opposite sides (as accurately as could be done) of the tips of 13 radicles, suspended in damp air, at a temperature of 65o - 66o F. The result was striking, for 9 out of these 13 radicles became plainly, and 1 very slightly, curved from the thick paper towards the side bearing the thin paper. In two of these cases the apex became completely hooked after two days; in four cases the deflection from the perpendicular and from the side bearing the thick paper, amounted in from two to four days to angles of 90o, 72o, 60o, and 49o, but in two other cases to only 18o and 15o. It should, however, be stated that in the [page 177]
case in which the deflection was 49o, the two squares had accidentally come into contact on one side of the apex, and thus formed a lateral gable; and the deflection was directed in part from this gable and in part from the thick paper. In three cases alone the radicles were not affected by the difference in thickness of the squares of paper attached to their tips, and consequently did not bend away from the side bearing the stiffer paper.
Zea mays: Sensitiveness of the apex of the Radicle to contact.--A large number of trials were made on this plant, as it was the only monocotyledon on which we experimented. An abstract of the results will suffice. In the first place, 22 germinating seeds were pinned to cork-lids without any object being attached to their radicles, some being exposed to a temperature of 65o - 66o F., and others to between 74o and 79o; and none of them became curved, though some were a little inclined to one side. A few were selected, which from having germinated on sand were crooked, but when suspended in damp air the terminal part grew straight downwards. This fact having been ascertained, little squares of the card-like paper were affixed with sh.e.l.lac, on several occasions, to the tips of 68 radicles. Of these the terminal growing part of 39 became within 24 h. conspicuously curved away from the attached squares and from the perpendicular; 13 out of the 39 forming hooks with their points directed towards the zenith, and 8 forming loops. Moreover, 7 other radicles out of the 68, were slightly and two doubtfully deflected from the cards. There remain 20 which were not affected; but 10 of these ought not to be counted; for one was diseased, two had their tips quite surrounded by sh.e.l.lac, and the squares on 7 had slipped so as to stand parallel to the apex, instead of obliquely [page 178]
on it. There were therefore only 10 out of the 68 which certainly were not acted on. Some of the radicles which were experimented on were young and short, most of them of moderate length, and two or three exceeded three inches in length. The curvature in the above cases occurred within 24 h., but it was often conspicuous within a much shorter period. For instance, the terminal growing part of one radicle was bent upwards into a rectangle in 8 h. 15 m., and of another in 9 h. On one occasion a hook was formed in 9 h. Six of the radicles in a jar containing nine seeds, which stood on a sand-bath, raised to a temperature varying from 76o to 82o F., became hooked, and a seventh formed a complete loop, when first looked at after 15 hours.
The accompanying figures of four germinating seeds (Fig. 69) show, firstly, a radicle (A) the apex of which has become so much bent away from the attached square as to form a hook. Secondly (B), a hook converted through the continued irritation of the card, aided perhaps by geotropism, into an almost complete circle or loop. The tip in the act of forming a loop generally rubs against the upper part of the radicle, and pushes off the attached square; the loop then contracts or closes, but never disappears; and the apex afterwards grows vertically downwards, being no longer irritated by any attached object. This frequently occurred, and is represented at C. The jar above mentioned with the six hooked radicles and another jar were kept for two additional days, for the sake of observing how the hooks would be modified. Most of them became converted into simple loops, like that figured at C; but in one case the apex did not rub against the upper part of the radicle and thus remove the card; and it consequently made, owing [page 179]
to the continued irritation from the card, two complete loops, that is, a helix of two spires; which afterwards became pressed closely together. Then geotropism prevailed and caused the apex to grow perpendicularly downwards.
In another case, shown at (D), the apex
Fig. 69. Zea mays: radicles excited to bend away from the little squares of card attached to one side of their tips.
in making a second turn or spire, pa.s.sed through the first loop, which was at first widely open, and in doing so knocked off the card; it then grew perpendicularly downwards, and thus tied itself into a knot, which soon became tight!
Secondary Radicles of Zea.--A short time after the first radicle has appeared, others protrude from the [page 180]
seed, but not laterally from the primary one. Ten of these secondary radicles, which were directed obliquely downwards, were experimented on with very small squares of card attached with sh.e.l.lac to the lower sides of their tips. If therefore the squares acted, the radicles would bend upwards in opposition to gravity. The jar stood (protected from light) on a sand-bath, which varied between 76o and 82o F. After only 5 h. one appeared to be a little deflected from the square, and after 20 h. formed a loop.
Four others were considerably curved from the squares after 20 h., and three of them became hooked, with their tips pointing to the zenith,--one after 29 h. and the two others after 44 h. By this latter time a sixth radicle had become bent at a right angle from the side bearing the square.
Thus altogether six out of the ten secondary radicles were acted on, four not being affected. There can, therefore, be no doubt that the tips of these secondary radicles are sensitive to slight contact, and that when thus excited they cause the upper part to bend from the touching object; but generally, as it appears, not in so short a time as in the case of the first-formed radicle.
SENSITIVENESS OF THE TIP OF THE RADICLE TO MOIST AIR.
Sachs made the interesting discovery, a few years ago, that the radicles of many seedling plants bend towards an adjoining damp surface.* We shall here endeavour to show that this peculiar form of sensitiveness resides in their tips. The movement is directly the reverse of that excited by the irritants. .h.i.therto considered, which cause the growing part of the
* "Arbeiten des Bot. Inst.i.tut., in Wurzburg," vol. i. 1872, p. 209.
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radicle to bend away from the source of irritation. In our experiments we followed Sachs" plan, and sieves with seeds germinating in damp sawdust were suspended so that the bottom was generally inclined at 40o with the horizon. If the radicles had been acted on solely by geotropism, they would have grown out of the bottom of the sieve perpendicularly downwards; but as they were attracted by the adjoining damp surface they bent towards it and were deflected 50o from the perpendicular. For the sake of ascertaining whether the tip or the whole growing part of the radicle was sensitive to the moist air, a length of from 1 to 2 mm. was coated in a certain number of cases with a mixture of olive-oil and lamp-black. This mixture was made in order to give consistence to the oil, so that a thick layer could be applied, which would exclude, at least to a large extent, the moist air, and would be easily visible. A greater number of experiments than those which were actually tried would have been necessary, had not it been clearly established that the tip of the radicle is the part which is sensitive to various other irritants.
[Phaseolus multiflorus.--Twenty-nine radicles, to which nothing had been done, growing out of a sieve, were observed at the same time with those which had their tips greased, and for an equal length of time. Of the 29, 24 curved themselves so as to come into close contact with the bottom of the sieve. The place of chief curvature was generally at a distance of 5 or 6 mm. from the apex. Eight radicles had their tips greased for a length of 2 mm., and two others for a length of 1 mm.; they were kept at a temperature of 15o - 16o C. After intervals of from 19 h. to 24 h. all were still vertically or almost vertically dependent, for some of them had moved towards the adjoining damp surface by about 10o. They had therefore not been acted on, or only slightly acted on, by the damper air on one side, although the whole upper part was freely exposed. After 48 h. three of these radicles became [page 182]
considerably curved towards the sieve; and the absence of curvature in some of the others might perhaps be accounted for by their not having grown very well. But it should be observed that during the first 19 h. to 24 h. all grew well; two of them having increased 2 and 3 mm. in length in 11 h.; five others increased 5 to 8 mm. in 19 h.; and two, which had been at first 4 and 6 mm. in length, increased in 24 h. to 15 and 20 mm.