Here, then, we have the rare opportunity of seeing a structure proper to a more advanced age, in the act of gradually encroaching on and replacing an earlier or embryological condition.
The genus Melilotus is closely allied to Trifolium, and the first leaf bears only a single leaflet, which at night rotates on its axis so as to present one lateral edge to the zenith. Hence it sleeps like the terminal leaflet of a mature plant, as was observed in 15 species, and wholly unlike the corresponding leaflet of Trifolium, which simply bends upwards. It is therefore a curious fact that in one of these 15 species, viz., M. Taurica (and in a lesser degree in two others), leaves arising from young shoots, produced on plants which had been cut down and kept in pots during the winter in the green-house, slept like the leaves of a Trifolium, whilst the leaves on the fully-grown branches on these same plants afterwards slept normally like those of a Melilotus. If young shoots rising from the ground may be considered as new individuals, partaking to a certain extent of the nature of seedlings, then the peculiar manner in which their leaves slept may be considered [page 416] as an embryological habit, probably the result of Melilotus being descended from some form which slept like a Trifolium. This view is partially supported by the leaves on old and young branches of another species, M. Messanensis (not included in the above 15 species), always sleeping like those of a Trifolium.
The first true leaf of Mimosa albida consists of a simple petiole, often bearing three pairs of leaflets, all of which are of nearly equal size and of the same shape: the second leaf differs widely from the first, and resembles that on a mature plant (see Fig. 159, p. 379), for it consists of two pinnae, each of which bears two pairs of leaflets, of which the inner basal one is very small. But at the base of each pinna there is a pair of minute points, evidently rudiments of leaflets, for they are of unequal sizes, like the two succeeding leaflets. These rudiments are in one sense embryological, for they exist only during the youth of the leaf, falling off and disappearing as soon as it is fully grown.
With Desmodium gyrans the two lateral leaflets are very much smaller than the corresponding leaflets in most of the species in this large genus; they vary also in position and size; one or both are sometimes absent; and they do not sleep like the fully-developed leaflets. They may therefore be considered as almost rudimentary; and in accordance with the general principles of embryology, they ought to be more constantly and fully developed on very young than on old plants. But this is not the case, for they were quite absent on some young seedlings, and did not appear until from 10 to 20 leaves had been formed. This fact leads to the suspicion that D. gyrans is descended through a unifoliate form (of which some exist) from a trifoliate species; and that the little lateral leaflets reappear through reversion. However this may be, [page 417] the interesting fact of the pulvini or organs of movement of these little leaflets, not having been reduced nearly so much as their blades--taking the large terminal leaflet as the standard of comparison--gives us probably the proximate cause of their extraordinary power of gyration. [page 418]
CHAPTER VIII.
MODIFIED CIRCUMNUTATION: MOVEMENTS EXCITED BY LIGHT.
Distinction between heliotropism and the effects of light on the periodicity of the movements of leaves--Heliotropic movements of Beta, Solanum, Zea, and Avena--Heliotropic movements towards an obscure light in Apios, Brassica, Phalaris, Tropaeolum, and Cassia--Apheliotropic movements of tendrils of Bignonia--Of flower-peduncles of Cyclamen--Burying of the pods--Heliotropism and apheliotropism modified forms of circumnutation-- Steps by which one movement is converted into the other-- Transversal-heliotropismus or diaheliotropism influenced by epinasty, the weight of the part and apogeotropism--Apogeotropism overcome during the middle of the day by diaheliotropism--Effects of the weight of the blades of cotyledons--So called diurnal sleep--Chlorophyll injured by intense light--Movements to avoid intense light
SACHS first clearly pointed out the important difference between the action of light in modifying the periodic movements of leaves, and in causing them to bend towards its source.* The latter, or heliotropic movements are determined by the direction of the light, whilst periodic movements are affected by changes in its intensity and not by its direction.