The Power of Movement in Plants, Charles Darwin [the reading list book TXT] 📗

in making a second turn or spire, passed 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

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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 shellac 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 hitherto considered, which cause the growing part of the * ‘Arbeiten des Bot. Institut., in W�rzburg,’ 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

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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.

The tips of 10 radicles, which likewise grew well, were coated with the grease for a length of only 1 mm., and now the result was somewhat different; for of these 4 curved themselves to the sieve in from 21 h. to 24h., whilst 6 did not do so. Five of the latter were observed for an additional day, and now all excepting one became curved to the sieve.

The tips of 5 radicles were cauterised with nitrate of silver, and about 1

mm. in length was thus destroyed. They were observed for periods varying between 11 h. and 24h., and were found to have grown well. One of them had curved until it came into contact with the sieve; another was curving towards it; whilst the remaining three were still vertically dependent. Of 7 not cauterised radicles observed at the same time, all had come into contact with the sieve.

The tips of 11 radicles were protected by moistened goldbeaters’ skin, which adheres closely, for a length varying from 1 � to 2 � mm. After 22 h.

to 24 h., 6 of these radicles were clearly bent towards or had come into contact with the sieve; 2 were slightly curved in this direction, and 3 not at all. All had grown well. Of 14 control specimens observed at the same time, all excepting one had closely approached the sieve. It appears from these cases that a cap of goldbeaters’ skin checks, though only to a slight degree, the bending of the radicles to an adjoining damp surface. Whether an extremely thin sheet of this substance when moistened allows moisture from the air to pass through it, we do not know. One case indicated that the caps were sometimes more efficient than appears from the above results; for a radicle, which after 23 h. had only slightly approached the sieve, had its cap (1 � mm. in length) removed, and during the next 15 � h. it curved itself abruptly towards the source of moisture, the chief seat of curvature being at a distance of 2 to 3 mm. from the apex.

Vicia faba.—The tips of 13 radicles were coated with the grease for a length of 2 mm.; and it should be remembered that with these radicles the seat of chief curvature is about

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4 or 5 mm. from the apex. Four of them were examined after 22h., three after 26 h., and six after 36 h., and none had been attracted towards the damp lower surface of the sieve. In another trial 7 radicles were similarly treated, and 5 of them still pointed perpendicularly downwards after 11 h., whilst 2 were a little curved towards the sieve; by an accident they were not subsequently observed. In both these trials the radicles grew well; 7

of them, which were at first from 4 to 11 mm. in length, were after 11 h.

between 7 and 16 mm.; 3 which were at first from 6 to 8 mm. after 26 h.

were 11.5 to 18 mm. in length; and lastly, 4 radicles which were at first 5

to 8 mm. after 46 h. were 18 to 23 mm. in length. The control or ungreased radicles were not invariably attracted towards the bottom of the sieve. But on one occasion 12 out of 13, which were observed for periods between 22 h.

and 36 h., were thus attracted. On two other occasions taken together, 38

out of 40 were similarly attracted. On another occasion only 7 out of 14

behaved in this manner, but after two more days the proportion of the curved increased to 17 out of 23. On a last occasion only 11 out of 20 were thus attracted. If we add up these numbers, we find that 78 out of 96 of the control specimens curved themselves towards the bottom of the sieve. Of the specimens with greased tips, 2 alone out of the 20 (but 7 of these were not observed for a sufficiently long time) thus curved themselves. We can, therefore, hardly doubt that the tip for a length of 2 mm. is the part which is sensitive to a moist atmosphere, 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 20o and 12o 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

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them not quite perpendicularly downwards, were coated with the black grease for a length of from 1 to 1 � mm. The sieves were inclined at 30o 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 10o and 35o; 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 1 � mm., they would not have been affected by the moist air and none would have become curved.

Triticum