It is not that they cease to circumnutate in darkness, for in all the many cases observed by us they continued to do so; but the normal order of their movements in relation to the alternations of day and night is much disturbed or quite annulled. This holds good with species the cotyledons of which rise or sink so much at night that they may be said to sleep, as well as with others which rise only a little. But different species are affected in very different degrees by changes in the light. [page 124]
[For instance, the cotyledons of Beta vulgaris, Solanum lycopersicum, Cerinthe major, and Lupinus luteus, when placed in darkness, moved down during the afternoon and early night, instead of rising as they would have done if they had been exposed to the light. All the individuals of the Solanum did not behave in the same manner, for the cotyledons of one circumnutated about the same spot between 2.30 and 10 P.M. The cotyledons of a seedling of Oxalis corniculata, which was feebly illuminated from above, moved downwards during the first morning in the normal manner, but on the second morning it moved upwards. The cotyledons of Lotus Jacoboeus were not affected by 4 h. of complete darkness, but when placed under a double skylight and thus feebly illuminated, they quite lost their periodical movements on the third morning. On the other hand, the cotyledons of Cucurbita ovifera moved in the normal manner during a whole day in darkness.
Seedlings of Githago segetum were feebly illuminated from above in the morning before their cotyledons had expanded, and they remained closed for the next 40 h. Other seedlings were placed in the dark after their cotyledons had opened in the morning and these did not begin to close until about 4 h. had elapsed. The cotyledons of Oxalis rosea sank vertically downwards after being left for 1 h. 20 m. in darkness; but those of some other species of Oxalis were not affected by several hours of darkness. The cotyledons of several species of Cassia are eminently susceptible to changes in the degree of light to which they are exposed: thus seedlings of an unnamed S. Brazilian species (a large and beautiful tree) were brought out of the hot-house and placed on a table in the middle of a room with two north-east and one north-west window, so that they were fairly well illuminated, though of course less so than in the hot-house, the day being moderately bright; and after 36 m. the cotyledons which had been horizontal rose up vertically and closed together as when asleep; after thus remaining on the table for 1 h. 13 m. they began to open. The cotyledons of young seedlings of another Brazilian species and of C. neglecta, treated in the same manner, behaved similarly, excepting that they did not rise up quite so much: they again became horizontal after about an hour.
Here is a more interesting case: seedlings of Cassia tora in two pots, which had stood for some time on the table in the room just described, had their cotyledons horizontal. One pot was now exposed for 2 h. to dull sunshine, and the cotyledons [page 125] remained horizontal; it was then brought back to the table, and after 50 m. the cotyledons had risen 68o above the horizon. The other pot was placed during the same 2 h. behind a screen in the room, where the light was very obscure, and the cotyledons rose 63o above the horizon; the pot was then replaced on the table, and after 50 m. the cotyledons had fallen 33o. These two pots with seedlings of the same age stood close together, and were exposed to exactly the same amount of light, yet the cotyledons in the one pot were rising, whilst those in the other pot were at the same time sinking. This fact illustrates in a striking manner that their movements are not governed by the actual amount, but by a change in the intensity or degree of the light. A similar experiment was tried with two sets of seedlings, both exposed to a dull light, but different in degree, and the result was the same. The movements of the cotyledons of this Cassia are, however, determined (as in many other cases) largely by habit or inheritance, independently of light; for seedlings which had been moderately illuminated during the day, were kept all night and on the following morning in complete darkness; yet the cotyledons were partially open in the morning and remained open in the dark for about 6 h.