Lastly, the wide distribution in the dicotyledonous series of plants with cotyledons which sleep. Reflecting on these several facts, our conclusion seems justified, that the nyctitropic movements of cotyledons, by which the blade is made to stand either vertically or almost vertically upwards or downwards at night, has been acquired, at least in most cases, for some special purpose; nor can we doubt that this purpose is the protection of the upper surface of the blade, and perhaps of the central bud or plumule, from radiation at night. [page 317]

CHAPTER VII.

MODIFIED CIRCUMNUTATION: NYCTITROPIC OR SLEEP MOVEMENTS OF LEAVES.

Conditions necessary for these movements--List of Genera and Families, which include sleeping plants--Description of the movements in the several Genera--Oxalis: leaflets folded at night--Averrhoa: rapid movements of the leaflets--Porlieria: leaflets close when plant kept very dry--Tropaeolum: leaves do not sleep unless well illuminated during day--Lupinus: various modes of sleeping--Melilotus: singular movements of terminal leaflet-- Trifolium--Desmodium: rudimentary lateral leaflets, movements of, not developed on young plants, state of their pulvini--Cassia: complex movements of the leaflets--Bauhinia: leaves folded at night--Mimosa pudica: compounded movements of leaves, effect of darkness--Mimosa albida, reduced leaflets of--Schrankia: downward movement of the pinnae--Marsilea: the only cryptogam known to sleep--Concluding remarks and summary--Nyctitropism consists of modified circumnutation, regulated by the alternations of light and darkness--Shape of first true leaves.

WE now come to the nyctitropic or sleep movements of leaves. It should be remembered that we confine this term to leaves which place their blades at night either in a vertical position or not more than 30o from the vertical,--that is, at least 60o above or beneath the horizon. In some few cases this is effected by the rotation of the blade, the petiole not being either raised or lowered to any considerable extent. The limit of 30o from the vertical is obviously an arbitrary one, and has been selected for reasons previously assigned, namely, that when the blade approaches the perpendicular as nearly as this, only half as much of the surface is exposed at night to the [page 318] zenith and to free radiation as when the blade is horizontal. Nevertheless, in a few instances, leaves which seem to be prevented by their structure from moving to so great an extent as 60o above or beneath the horizon, have been included amongst sleeping plants.

It should be premised that the nyctitropic movements of leaves are easily affected by the conditions to which the plants have been subjected. If the ground is kept too dry, the movements are much delayed or fail: according to Dassen,* even if the air is very dry the leaves of Impatiens and Malva are rendered motionless. Carl Kraus has also lately insisted** on the great influence which the quantity of water absorbed has on the periodic movements of leaves; and he believes that this cause chiefly determines the variable amount of sinking of the leaves of Polygonum convolvulus at night; and if so, their movements are not in our sense strictly nyctitropic. Plants in order to sleep must have been exposed to a proper temperature: Erythrina crista-galli, out of doors and nailed against a wall, seemed in fairly good health, but the leaflets did not sleep, whilst those on another plant kept in a warm greenhouse were all vertically dependent at night. In a kitchen-garden the leaflets of Phaseolus vulgaris did not sleep during the early part of the summer. Ch. Royer says,*** referring I suppose to the native plants in France, that they do not sleep when the temperature is below 5o C. or 41o F. In the case of several sleeping plants, viz., species of

* Dassen,'Tijdschrift vor. Naturlijke Gesch. en Physiologie,' 1837, vol. iv. p. 106. See also Ch. Royer on the importance of a proper state of turgescence of the cells, in 'Annal. des Sc. Nat.

Charles Darwin

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