The upward movement thus becomes unequal in rate, and is sometimes interrupted by stationary periods. Whenever irregular ellipses or loops are still formed, their longer axes are almost always directed in the line of gravity, in an analogous manner as occurred with heliotropic movements in reference to the light. As apogeotropism acts more and more energetically, ellipses or loops cease to be formed, and the course becomes at first strongly, and then less and less zigzag, and finally rectilinear. From this gradation in the nature of the movement, and more especially from all growing parts, which alone (except when pulvini are present) are acted on by apogeotropism, con- [page 508] tinually circumnutating, we may conclude that even a rectilinear course is merely an extremely modified form of circumnutation. It is remarkable that a stem or other organ which is highly sensitive to apogeotropism, and which has bowed itself rapidly upwards in a straight line, is often carried beyond the vertical, as if by momentum. It then bends a little backwards to a point round which it finally circumnutates. Two instances of this were observed with the hypocotyls of Beta vulgaris, one of which is shown in Fig. 183, and two other instances with the hypocotyls of Brassica. This momentum-like movement probably results from the accumulated effects of apogeotropism. For the sake of observing how long such after-effects lasted, a pot with seedlings of Beta was laid on its side in the dark, and the hypocotyls in 3 h. 15 m. became highly inclined. The pot, still in the dark, was then placed upright, and the movements of the two hypocotyls were traced; one continued to bend in its former direction, now in opposition to apogeotropism, for about 37 m., perhaps for 48 m.; but after 61 m. it moved in an opposite direction. The other hypocotyl continued to move in its former course, after being placed upright, for at least 37 m.

Different species and different parts of the same species are acted on by apogeotropism in very different degrees. Young seedlings, most of which circumnutate quickly and largely, bend upwards and become vertical in much less time than do any older plants observed by us; but whether this is due to their greater sensitiveness to apogeotropism, or merely to their greater flexibility we do not know. A hypocotyl of Beta traversed an angle of 109o in 3 h. 8 m., and a cotyledon of Phalaris an angle of 130o in 4 h. 30 m. On the other hand, the stem of a herbaceous [page 509] Verbena rose 90o in about 24 h.; that of Rubus 67o, in 70 h.; that of Cytisus 70o, in 72 h.; that of a young American Oak only 37o, in 72 h. The stem of a young Cyperus alternifolius rose only 11o in 96 h.; the bending being confined to near its base. Though the sheath-like cotyledons of Phalaris are so extremely sensitive to apogeotropism, the first true leaves which protrude from them exhibited only a trace of this action. Two fronds of a fern, Nephrodium molle, both of them young and one with the tip still inwardly curled, were kept in a horizontal position for 46 h., and during this time they rose so little that it was doubtful whether there was any true apogeotropic movement.

The most curious case known to us of a difference in sensitiveness to gravitation, and consequently of movement, in different parts of the same organ, is that offered by the petioles of the cotyledons of Ipomoea leptophylla. The basal part for a short length where united to the undeveloped hypocotyl and radicle is strongly geotropic, whilst the whole upper part is strongly apogeotropic. But a portion near the blades of the cotyledons is after a time acted on by epinasty and curves downwards, for the sake of emerging in the form of an arch from the ground; it subsequently straightens itself, and is then again acted on by apogeotropism.

A branch of Cucurbita ovifera, placed horizontally, moved upwards during 7 h. in a straight line, until it stood at 40o above the horizon; it then began to circumnutate, as if owing to its trailing nature it had no tendency to rise any higher.

Charles Darwin

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