Another upright branch was secured to a stick, close to the base of a tendril, and the pot was then laid horizontally in the dark. In this position the tendril circumnutated and made [page 510] several large ellipses during 14 h., as it likewise did on the following day; but during this whole time it was not in the least affected by apogeotropism. On the other hand, when branches of another Cucurbitaceous plant, Echinocytis lobata, were fixed in the dark so that the tendrils depended beneath the horizon, these began immediately to bend upwards, and whilst thus moving they ceased to circumnutate in any plain manner; but as soon as they had become horizontal they recommenced to revolve conspicuously.* The tendrils of Passiflora gracilis are likewise apogeotropic. Two branches were tied down so that their tendrils pointed many degrees beneath the horizon. One was observed for 8 h., during which time it rose, describing two circles, one above the other. The other tendril rose in a moderately straight line during the first 4 h., making however one small loop in its course; it then stood at about 45o above the horizon, where it circumnutated during the remaining 8 h. of observation.

A part or organ which whilst young is extremely sensitive to apogeotropism ceases to be so as it grows old; and it is remarkable, as showing the independence of this sensitiveness and of the circumnutating movement, that the latter sometimes continues for a time after all power of bending from the centre of the earth has been lost. Thus a seedling Orange bearing only 3 young leaves, with a rather stiff stem, did not curve in the least upwards during 24 h. whilst extended horizontally; yet it circumnutated all the time over a small space. The hypocotyl of a young seedling of Cassia tora, similarly placed, became vertical in 12 h.; that of an older seedling, 1 1/4 inch in height,

* For details see 'The Movements and Habits of Climbing Plants,' 1875, p. 131. [page 511]

became so in 28 h.; and that of another still older one, 1 ½ inch in height, remained horizontal during two days, but distinctly circumnutated during this whole time.

When the cotyledons of Phalaris or Avena are laid horizontally, the uppermost part first bends upwards, and then the lower part; consequently, after the lower part has become much curved upwards, the upper part is compelled to curve backwards in an opposite direction, in order to straighten itself and to stand vertically; and this subsequent straightening process is likewise due to apogeotropism. The upper part of 8 young cotyledons of Phalaris were made rigid by being cemented to thin glass rods, so that this part could not bend in the least; nevertheless, the basal part was not prevented from curving upward. A stem or other organ which bends upwards through apogeotropism exerts considerable force; its own weight, which has of course to be lifted, was sufficient in almost every instance to cause the part at first to bend a little downwards; but the downward course was often rendered oblique by the simultaneous circumnutating movement. The cotyledons of Avena placed horizontally, besides lifting their own weight, were able to furrow the soft sand above them, so as to leave little crescentic open spaces on the lower sides of their bases; and this is a remarkable proof of the force exerted.

As the tips of the cotyledons of Phalaris and Avena bend upwards through the action of apogeotropism before the basal part, and as these same tips when excited by a lateral light transmit some influence to the lower part, causing it to bend, we thought that the same rule might hold good with apogeotropism. Consequently, the tips of 7 cotyledons of Phalaris were [page 512] cut off for a length in three cases of .2 inch and in the four other cases of .14, .12, .1, and .07 inch. But these cotyledons, after being extended horizontally, bowed themselves upwards as effectually as the unmutilated specimens in the same pots, showing that sensitiveness to gravitation is not confined to their tips.

Charles Darwin

All Pages of This Book