cotyledons, and the movement of the whole seedling was traced during two days. The circumnutation of the hypocotyl is quite insignificant compared with that of the cotyledons. These rise up vertically at night and come into close contact; so that they may be said to sleep. This seedling was so old that a very small true leaf had been developed, which at night was completely hidden by the closed cotyledons. On Sept.
24th, between 8 A.M. and 5 P.M., the cotyledons moved five times up and five times down; they therefore described five irregular ellipses in the course of the 9 h. The great nocturnal rise commenced about 4.30 P.M.
[1] Seeds of this plant, which grew near the sea-side, were sent to us by Fritz Müller from S. Brazil. The seedlings did not flourish or flower well with us; they were sent to Kew, and were pronounced not to be distinguishable from C. tora.
Fig. 25. Cassia tora: conjoint circumnutation of cotyledons and hypocotyl, traced on vertical glass, from 7.10 A.M. Sept. 25th to 7.30 A.M. 26th. Figure here given reduced to one-half of original scale.
On the following morning (Sept. 25th) the movement of the same cotyledon was again traced in the same manner during 24 h.; and a copy of the tracing is here given (Fig. 25). The morning was cold, and the window had been accidentally left open for a short time, which must have chilled the plant; and this probably prevented it from moving quite as freely as on the previous day; for it rose only four and sank only four times during the day, one of the oscillations being very small. At 7.10 A.M., when the first dot was made, the cotyledons were not fully open or awake; they continued to open till about 9 A.M., by which time they had sunk a little beneath the horizon: by 9.30 A.M.
they had risen, and then they oscillated up and down; but the upward and downward lines never quite coincided. At about 4.30 P.M. the great nocturnal rise commenced. At 7 A.M. on the following morning (Sept.
26th) they occupied nearly the same level as on the previous morning, as shown in the diagram: they then began to open or sink in the usual manner. The diagram leads to the belief that the great periodical daily rise and fall does not differ essentially, excepting in amplitude, from the oscillations during the middle of the day.
Lotus Jacoboeus (Leguminosae).—The cotyledons of this plant, after the few first days of their life, rise so as to stand almost, though rarely quite, vertically at night. They continue to act in this manner for a long time even after the development of some of the true leaves. With seedlings, 3 inches in height, and bearing five or six leaves, they rose at night about 45°. They continued to act thus for about an additional fortnight. Subsequently they remained horizontal at night, though still green and at last dropped off. Their rising at night so as to stand almost vertically appears to depend largely on temperature; for when the seedlings were kept in a cool house, though they still continued to grow, the cotyledons did not become vertical at night. It is remarkable that the cotyledons do not generally rise at night to any conspicuous extent during the first four or five days after germination; but the period was extremely variable with seedlings kept under the same conditions; and many were observed. Glass filaments with minute triangles of paper were fixed to the cotyledons (1½ mm. in breadth) of two seedlings, only 24 h. old, and the hypocotyl was secured to a stick; their movements greatly magnified were traced, and they certainly circumnutated the whole time on a small scale, but they did not exhibit any distinct nocturnal and diurnal movement. The hypocotyls, when left free, circumnutated over a large space.
Another and much older seedling, bearing a half-developed leaf, had its movements traced in a similar manner during the three first days and nights of June; but seedlings at this age appear to be very sensitive to a deficiency of light; they were observed under a rather dim skylight, at a temperature of between 16° to 17½° C.’ and apparently, in consequence of these conditions, the great daily movement of the cotyledons ceased on the third day. During the first two days they began rising in the early afternoon in a nearly straight line, until between 6 and 7 P.M., when they stood vertically. During the latter part of the night, or more probably in the early morning, they began to fall or open, so that by 6.45 A.M. they stood fully expanded and horizontal. They continued to fall slowly for some time, and during the second day described a single small ellipse, between 9 A.M. and 2 P.M., in addition to the great diurnal movement. The course pursued during the whole 24 h. was far less complex than in the foregoing case of Cassia. On the third morning they fell very much, and then circumnutated on a small scale round the same spot; by 8.20 P.M. they showed no tendency to rise at night. Nor did the cotyledons of any of the many other seedlings in the same pot rise; and so it was on the following night of June 5th. The pot was then taken back into the hot-house, where it was exposed to the sun, and on the succeeding night all the cotyledons rose again to a high angle, but did not stand quite vertically. On each of the above days the line representing the great nocturnal rise did not coincide with that of the great diurnal fall, so that narrow ellipses were described, as is the usual rule with circumnutating organs. The cotyledons are provided with a pulvinus, and its development will hereafter be described.
Mimosa pudica (Leguminosae).—The cotyledons rise up vertically at night, so as to close together. Two seedlings were observed in the greenhouse (temp. 16° to 17° C. or 63° to 65° F.). Their hypocotyls were secured to sticks, and glass filaments bearing little triangles of paper were affixed to the cotyledons of both. Their movements were traced on a vertical glass during 24 h. on November 13th. The pot had stood for some time in the same position, and they were chiefly illuminated through the glass-roof. The cotyledons of one of these seedlings moved downward in the morning till 11.30 A.M., and then rose, moving rapidly in the evening until they stood vertically, so that in this case there was simply a single great daily fall and rise. The other seedling behaved rather differently, for it fell in the morning until 11.30 A.M., and then rose, but after 12.10 P.M. again fell; and the great evening rise did not begin until 1.22 P.M. On the following morning this cotyledon had fallen greatly from its vertical position by 8.15 A.M. Two other seedlings (one seven and the other eight days old) had been previously observed under unfavourable circumstances, for they had been brought into a room and placed before a north-east window, where the temperature was between only 56° and 57° F. They had, moreover, to be protected from lateral light, and perhaps were not sufficiently illuminated. Under these circumstances the cotyledons moved simply downwards from 7 A.M. till 2 P.M., after which hour and during a large part of the night they continued to rise. Between 7 and 8 A.M. on the following morning they fell again; but on this second and likewise on the third day the movements became irregular, and between 3 and 10.30 P.M. they circumnutated to a small extent about the same spot; but they did not rise at night. Nevertheless, on the following night they rose as usual.
Cytisus fragrans (Leguminosae).—Only a few observations were made on this plant. The hypocotyl circumnutated to a considerable extent, but in a simple manner—namely, for two hours in one direction, and then much more slowly back again in a zigzag course, almost parallel to the first line, and beyond the starting-point. It moved in the same direction all night, but next morning began to return. The cotyledons continually move both up and down and laterally; but they do not rise up at night in a conspicuous manner.
Lupinus luteus (Leguminosae).—Seedlings of this plant were observed because the cotyledons are so thick (about.08 of an inch) that it seemed unlikely that they would move. Our observations were not very successful, as the seedlings are strongly heliotropic, and their circumnutation could not be accurately observed near a north-east window, although they had been kept during the previous day in the same position. A seedling was then placed in darkness with the hypocotyl secured to a stick; both cotyledons rose a little at first, and then fell during the rest of the day; in the evening between 5 and 6 P.M.
they moved very slowly; during the night one continued to fall and the other rose, though only a little. The tracing was not much magnified, and as the lines were plainly zigzag, the cotyledons must have moved a little laterally, that is, they must have circumnutated.
The hypocotyl is rather thick, about.12 of inch; nevertheless it circumnutated in a complex course, though to a small extent. The movement of an old seedling with two true leaves partially developed, was observed in the dark. As the movement was magnified about 100 times it is not trustworthy and is not given; but there could be no doubt that the hypocotyl moved in all directions during the day, changing its course 19 times. The extreme actual distance from side to side through which the upper part of the hypocotyl passed in the course of 14½ hours was only 1/60 of an inch; it sometimes travelled at the rate of 1/50 of an inch in an hour.
Cucurbita ovifera (Cucurbitaceæ).—Radicle: a seed which had germinated on damp sand was fixed so that the slightly curved radicle, which was only.07 inch in length, stood almost vertically upwards, in which position geotropism would act at first with little power. A filament was attached near to its base, and projected at about an angle of 45° above the horizon. The general course followed during the 11 hours of observation and during the following night is shown in the accompanying diagram (Fig. 26), and was plainly due to geotropism; but it was also clear that the radicle circumnutated. By the next morning the tip had curved so much downwards that the filament, instead of projecting at 45° above the horizon, was nearly horizontal. Another germinating seed was turned upside down and covered with damp sand; and a filament was fastened to the radicle so as to project at an angle of about 50° above the horizon; this radicle was.35 of an inch in length and a little curved. The course pursued was mainly governed, as in the last case, by geotropism, but the line traced during 12 hours and magnified as before was more strongly zigzag, again showing circumnutation.
Fig. 26. Cucurbita ovifera: course followed by a radicle in bending geotropically downwards, traced on a horizontal glass, between 11.25 A.M. and 10.25 P.M.; the direction during the night is indicated by the broken line. Movement of bead magnified 14 times.
Four radicles were allowed to grow downwards over plates of smoked glass, inclined at 70° to the horizon, under the same conditions as in the cases of Æsculus, Phaseolus, and Vicia. Facsimiles are here given (Fig. 27) of two of these tracks; and a third short one was almost as plainly serpentine as that at A. It was also manifest by a greater or less amount of soot having been swept off the glasses, that the tips had pressed alternately with greater and less force on them. There must, therefore, have been movement in at least two planes at right angles to one another. These radicles were so delicate that they rarely had the power to sweep the glasses quite clean. One of them had developed some lateral or secondary rootlets, which projected a few degrees beneath the horizon; and it is an important fact that three of them left distinctly serpentine tracks on the smoked surface, showing beyond doubt that they had circumnutated like the main or primary radicle. But the tracks were so slight that they could not be traced and copied after the smoked surface had been varnished.
Fig. 27. Cucurbita ovifera: tracks left by tips of radicles in growing downwards over smoked glass-plates, inclined at 70° to the horizon.
Fig. 28. Cucurbita ovifera: circumnutation of arched hypocotyl at a very early age, traced in darkness on a horizontal glass, from 8 A.M.
to 10.20 A.M. on the following day. The movement of the bead magnified 20 times, here reduced to one-half of original scale.
Fig. 29. Cucurbita ovifera: circumnutation of straight and vertical hypocotyl, with filament fastened transversely across its upper end, traced in darkness on a horizontal glass, from 8.30 A.M. to 8.30 P.M.
The movement of the terminal bead originally magnified about 18 times, here only 4½ times.
Hypocotyl.—A seed lying on damp sand was firmly fixed by two crossed wires and by its own growing radicle. The cotyledons were still enclosed within the seed-coats; and the short hypocotyl, between the summit of the radicle and the cotyledons, was as yet only slightly arched. A filament (.85 of inch in length) was attached at an angle of 35° above the horizon to the side of the arch adjoining the cotyledons.
This part would ultimately form the upper end of the hypocotyl, after it had grown straight and vertical. Had the seed been properly planted, the hypocotyl at this stage of growth would have been deeply buried beneath the surface. The course followed by the bead of the filament is shown in Fig. 28. The chief lines of movement from left to right in the figure were parallel to the plane of the two united cotyledons and of the flattened seed; and this movement would aid in dragging them out of the seed-coats, which are held down by a special structure hereafter to be described. The movement at right angles to the above lines was due to the arched hypocotyl becoming more arched as it increased in height.
The foregoing observations apply to the leg of the arch next to the cotyledons, but the other leg adjoining the radicle likewise circumnutated at an equally early age.
The movement of the same hypocotyl after it had become straight and vertical, but with the cotyledons only partially expanded, is shown in Fig. 29. The course pursued during 12 h. apparently represents four and a half ellipses or ovals, with the longer axis of the first at nearly right angles to that of the others. The longer axes of all were oblique to a line joining the opposite cotyledons. The actual extreme distance from side to side over which the summit of the tall hypocotyl passed in the course of 12 h. was.28 of an inch. The original figure was traced on a large scale, and from the obliquity of the line of view the outer parts of the diagram are much exaggerated.
Cotyledons.—On two occasions the movements of the cotyledons were traced on a vertical glass, and as the ascending and descending lines did not quite coincide, very narrow ellipses were formed; they therefore circumnutated. Whilst young they rise vertically up at night, but their tips always remain reflexed; on the following morning they sink down again. With a seedling kept in complete darkness they moved in the same manner, for they sank from 8.45 A.M. to 4.30 P.M.; they then began to rise and remained close together until 10 P.M., when they were last observed. At 7 A.M. on the following morning they were as much expanded as at any hour on the previous day. The cotyledons of another young seedling, exposed to the light, were fully open for the first time on a certain day, but were found completely closed at 7 A.M.
on the following morning. They soon began to expand again, and continued doing so till about 5 P.M.; they then began to rise, and by 10.30 P.M. stood vertically and were almost closed. At 7 A.M. on the third morning they were nearly vertical, and again expanded during the day; on the fourth morning they were not closed, yet they opened a little in the course of the day and rose a little on the following night. By this time a minute true leaf had become developed. Another seedling, still older, bearing a well-developed leaf, had a sharp rigid filament affixed to one of its cotyledons (85 mm. in length), which recorded its own movements on a revolving drum with smoked paper. The observations were made in the hot-house, where the plant had lived, so that there was no change in temperature or light. The record commenced at 11 A.M. on February 18th; and from this hour till 3 P.M. the cotyledon fell; it then rose rapidly till 9 P.M., then very gradually till 3 A.M. February 19th, after which hour it sank gradually till 4.30 P.M.; but the downward movement was interrupted by one slight rise or oscillation about 1.30 P.M. After 4.30 P.M. (19th) the cotyledon rose till 1 A.M. (in the night of February 20th) and then sank very gradually till 9.30 A.M., when our observations ceased. The amount of movement was greater on the 18th than on the 19th or on the morning of the 20th.
Cucurbita aurantia.—An arched hypocotyl was found buried a little beneath the surface of the soil; and in order to prevent it straightening itself quickly, when relieved from the surrounding pressure of the soil, the two legs of the arch were tied together. The seed was then lightly covered with loose damp earth. A filament with a bead at the end was affixed to the basal leg, the movements of which were observed during two days in the usual manner. On the first day the arch moved in a zigzag line towards the side of the basal leg. On the next day, by which time the dependent cotyledons had been dragged above the surface of the soil, the tied arch changed its course greatly nine times in the course of 14½ h. It swept a large, extremely irregular, circular figure, returning at night to nearly the same spot whence it had started early in the morning. The line was so strongly zigzag that it apparently represented five ellipses, with their longer axes pointing in various directions. With respect to the periodical movements of the cotyledons, those of several young seedlings formed together at 4 P.M. an angle of about 60°, and at 10 P.M. their lower parts stood vertically and were in contact; their tips, however, as is usual in the genus, were permanently reflexed. These cotyledons, at 7 A.M. on the following morning, were again well expanded.
Lagenaria vulgaris (var. miniature Bottle-gourd) (Cucurbitaceæ).—A seedling opened its cotyledons, the movements of which were alone observed, slightly on June 27th and closed them at night: next day, at noon (28th), they included an angle of 53°, and at 10 P.M. they were in close contact, so that each had risen 26½°. At noon, on the 29th, they included an angle of 118°, and at 10 P.M. an angle of 54°, so each had risen 32°. On the following day they were still more open, and the nocturnal rise was greater, but the angles were not measured. Two other seedlings were observed, and behaved during three days in a closely similar manner. The cotyledons, therefore, open more and more on each succeeding day, and rise each night about 30°; consequently during the first two nights of their life they stand vertically and come into contact.
Fig. 30. Lagenaria vulgaris: circumnutation of a cotyledon, 1½ inch in length, apex only 4 3/4 inches from the vertical glass, on which its movements were traced from 7.35 A.M. July 11th to 9.5 A.M. on the 14th.
Figure here given reduced to one-third of original scale.
In order to ascertain more accurately the nature of these movements, the hypocotyl of a seedling, with its cotyledons well expanded, was secured to a little stick, and a filament with triangles of paper was affixed to one of the cotyledons. The observations were made under a rather dim skylight, and the temperature during the whole time was between 17½° to 18° C. (63° to 65° F.). Had the temperature been higher and the light brighter, the movements would probably have been greater.
On July 11th (see Fig. 30), the cotyledon fell from 7.35 A.M. till 10 A.M.; it then rose (rapidly after 4 P.M.) till it stood quite vertically at 8.40 P.M. During the early morning of the next day (12th) it fell, and continued to fall till 8 A.M., after which hour it rose, then fell, and again rose, so that by 10.35 P.M. it stood much higher than it did in the morning, but was not vertical as on the preceding night. During the following early morning and whole day (13th) it fell and circumnutated, but had not risen when observed late in the evening; and this was probably due to the deficiency of heat or light, or of both. We thus see that the cotyledons became more widely open at noon on each succeeding day; and that they rose considerably each night, though not acquiring a vertical position, except during the first two nights.
Cucumis dudaim (Cucurbitaceæ).—Two seedlings had opened their cotyledons for the first time during the day,—one to the extent of 90° and the other rather more; they remained in nearly the same position until 10.40 P.M.; but by 7 A.M. on the following morning the one which had been previously open to the extent of 90° had its cotyledons vertical and completely shut; the other seedling had them nearly shut. Later in the morning they opened in the ordinary manner.
It appears therefore that the cotyledons of this plant close and open at somewhat different periods from those of the foregoing species of the allied genera of Cucurbita and Lagenaria.
Fig. 31. Opuntia basilaris: conjoint circumnutation of hypocotyl and cotyledon; filament fixed longitudinally to cotyledon, and movement traced during 66 h. on horizontal glass. Movement of the terminal bead magnified about 30 times, here reduced to one-third scale. Seedling kept in hot-house, feebly illuminated from above.
Opuntia basilaris (Cacteæ).—A seedling was carefully observed, because, considering its appearance and the nature of the mature plant, it seemed very unlikely that either the hypocotyl or cotyledons would circumnutate to an appreciable extent. The cotyledons were well developed, being.9 of an inch in length,.22 in breadth, and.15 in thickness. The almost cylindrical hypocotyl, now bearing a minute spinous bud on its summit, was only.45 of an inch in height, and.19 in diameter. The tracing (Fig. 31) shows the combined movement of the hypocotyl and of one of the cotyledons, from 4.45 P.M. on May 28th to 11 A.M. on the 31st. On the 29th a nearly perfect ellipse was completed. On the 30th the hypocotyl moved, from some unknown cause, in the same general direction in a zigzag line; but between 4.30 and 10 P.M. almost completed a second small ellipse. The cotyledons move only a little up and down: thus at 10.15 P.M. they stood only 10° higher than at noon. The chief seat of movement therefore, at least when the cotyledons are rather old as in the present case, lies in the hypocotyl. The ellipse described on the 29th had its longer axis directed at nearly right angles to a line joining the two cotyledons.
The actual amount of movement of the bead at the end of the filament was, as far as could be ascertained, about.14 of an inch.
Fig. 32. Helianthus annuus: circumnutation of hypocotyl, with filament fixed across its summit, traced on a horizontal glass in darkness, from 8.45 A.M. to 10.45 P.M., and for an hour on following morning. Movement of bead magnified 21 times, here reduced to one-half of original scale.
Helianthus annuus (Compositæ).—The upper part of the hypocotyl moved during the day-time in the course shown in the annexed figure (Fig.
32). As the line runs in various directions, crossing itself several times, the movement may be considered as one of circumnutation. The extreme actual distance travelled was at least.1 of an inch. The movements of the cotyledons of two seedlings were observed; one facing a north-east window, and the other so feebly illuminated from above us as to be almost in darkness. They continued to sink till about noon, when they began to rise; but between 5 and 7 or 8 P.M. they either sank a little, or moved laterally, and then again began to rise. At 7 A.M.
on the following morning those on the plant before the north-east window had opened so little that they stood at an angle of 73° above the horizon, and were not observed any longer. Those on the seedling which had been kept in almost complete darkness, sank during the whole day, without rising about mid-day, but rose during the night. On the third and fourth days they continued sinking without any alternate ascending movement; and this, no doubt, was due to the absence of light.
Primula Sinensis (Primulaceae).—A seedling was placed with the two cotyledons parallel to a north-east window on a day when the light was nearly uniform, and a filament was affixed to one of them. From observations subsequently made on another seedling with the stem secured to a stick, the greater part of the movement shown in the annexed figure (Fig. 33), must have been that of the hypocotyl, though the cotyledons certainly move up and down to a certain extent both during the day and night. The movements of the same seedling were traced on the following day with nearly the same result; and there can be no doubt about the circumnutation of the hypocotyl.
Fig. 33. Primula Sinensis: conjoint circumnutation of hypocotyl and cotyledon, traced on vertical glass, from 8.40 A.M. to 10.45 P.M.
Movements of bead magnified about 26 times.
Cyclamen Persicum (Primulaceae).—This plant is generally supposed to produce only a single cotyledon, but Dr. H. Gressner[2] has shown that a second one is developed after a long interval of time. The hypocotyl is converted into a globular corm, even before the first cotyledon has broken through the ground with its blade closely enfolded and with its petiole in the form of an arch, like the arched hypocotyl or epicotyl of any ordinary dicotyledonous plant. A glass filament was affixed to a cotyledon,.55 of an inch in height, the petiole of which had straightened itself and stood nearly vertical, but with the blade not as yet fully expanded. Its movements were traced during 24½ h. on a horizontal glass, magnified 50 times; and in this interval it described two irregular small circles; it therefore circumnutates, though on an extremely small scale.
Fig. 34. Stapelia sarpedon: circumnutation of hypocotyl, illuminated from above, traced on horizontal glass, from 6.45 A.M. June 26th to 8.45 A.M. 28th. Temp. 23–24° C. Movement of bead magnified 21 times.
Stapelia sarpedon (Asclepiadeae).—This plant, when mature, resembles a cactus. The flattened hypocotyl is fleshy, enlarged in the upper part, and bears two rudimentary cotyledons. It breaks through the ground in an arched form, with the rudimentary cotyledons closed or in contact. A filament was affixed almost vertically to the hypocotyl of a seedling half an inch high; and its movements were traced during 50 h. on a horizontal glass (Fig. 34).
From some unknown cause it bowed itself to one side, and as this was effected by a zigzag course, it probably circumnutated; but with hardly any other seedling observed by us was this movement so obscurely shown.
Ipomœa caerulea vel Pharbitis nil (Convolvulaceae).—Seedlings of this plant were observed because it is a twiner, the upper internodes of which circumnutate conspicuously; but like other twining plants, the first few internodes which rise above the ground are stiff enough to support themselves, and therefore do not circumnutate in any plainly recognisable manner.[3] In this particular instance the fifth internode (including the hypocotyl) was the first which plainly circumnutated and twined round a stick. We therefore wished to learn whether circumnutation could be observed in the hypocotyl if carefully observed in our usual manner. Two seedlings were kept in the dark with filaments fixed to the upper part of their hypocotyls; but from circumstances not worth explaining their movements were traced for only a short time. One moved thrice forwards and twice backwards in nearly opposite directions, in the course of 3 h. 15 m.; and the other twice forwards and twice backwards in 2 h. 22 m. The hypocotyl therefore circumnutated at a remarkably rapid rate. It may here be added that a filament was affixed transversely to the summit of the second internode above the cotyledons of a little plant 3½ inches in height; and its movements were traced on a horizontal glass. It circumnutated, and the actual distance travelled from side to side was a quarter of an inch, which was too small an amount to be perceived without the aid of marks.
The movements of the cotyledons are interesting from their complexity and rapidity, and in some other respects. The hypocotyl (2 inches high) of a vigorous seedling was secured to a stick, and a filament with triangles of paper was affixed to one of the cotyledons. The plant was kept all day in the hot-house, and at 4.20 P.M. (June 20th) was placed under a skylight in the house, and observed occasionally during the evening and night. It fell in a slightly zigzag line to a moderate extent from 4.20 P.M. till 10.15 P.M. When looked at shortly after midnight (12.30 P.M.) it had risen a very little, and considerably by 3.45 A.M. When again looked at, at 6.10 A.M. (21st), it had fallen largely. A new tracing was now begun (see Fig. 35), and soon afterwards, at 6.42 A.M., the cotyledon had risen a little. During the forenoon it was observed about every hour; but between 12.30 and 6 P.M.
every half-hour. If the observations had been made at these short intervals during the whole day, the figure would have been too intricate to have been copied. As it was, the cotyledon moved up and P.M.) thirteen times.
Fig. 35. Ipomœa caerulea: circumnutation of cotyledon, traced on vertical glass, from 6.10 A.M. June 21st to 6.45 A.M. 22nd. Cotyledon with petiole 1.6 inch in length, apex of blade 4.1 inch from the vertical glass; so movement not greatly magnified; temp. 20° C.
The cotyledons of this seedling sank downwards during both evenings and the early part of the night, but rose during the latter part. As this is an unusual movement, the cotyledons of twelve other seedlings were observed; they stood almost or quite horizontally at mid-day, and at 10 P.M. were all declined at various angles. The most usual angle was between 30° and 35°; but three stood at about 50° and one at even 70° beneath the horizon. The blades of all these cotyledons had attained almost their full size, viz. from 1 to 1½ inches in length, measured along their midribs. It is a remarkable fact that whilst young—that is, when less than half an inch in length, measured in the same manner—they do not sink downwards in the evening. Therefore their weight, which is considerable when almost fully developed, probably came into play in originally determining the downward movement. The periodicity of this movement is much influenced by the degree of light to which the seedlings have been exposed during the day; for three kept in an obscure place began to sink about noon, instead of late in the evening; and those of another seedling were almost paralysed by having been similarly kept during two whole days. The cotyledons of several other species of Ipomœa likewise sink downwards late in the evening.
Cerinthe major (Boragineae).—The circumnutation of the hypocotyl of a young seedling with the cotyledons hardly expanded, is shown in the annexed figure (Fig. 36), which apparently represents four or five irregular ellipses, described in the course of a little over 12 hours.
Two older seedlings were similarly observed, excepting that one of them was kept in the dark; their hypocotyls also circumnutated, but in a more simple manner. The cotyledons on a seedling exposed to the light fell from the early morning until a little after noon, and then continued to rise until 10.30 P.M. or later. The cotyledons of this same seedling acted in the same general manner during the two following days. It had previously been tried in the dark, and after being thus kept for only 1 h. 40 m. the cotyledons began at 4.30 P.M. to sink, instead of continuing to rise till late at night.
Fig. 36. Cerinthe major: circumnutation of hypocotyl, with filament fixed across its summit, illuminated from above, traced on horizontal glass, from 9.26 A.M. to 9.53 P.M. on Oct. 25th. Movement of the bead magnified 30 times, here reduced to one-third of original scale.
Nolana prostrata (Nolaneae).—The movements were not traced, but a pot with seedlings, which had been kept in the dark for an hour, was placed under the microscope, with the micrometer eye-piece so adjusted that each division equalled 1/500th of an inch. The apex of one of the cotyledons crossed rather obliquely four divisions in 13 minutes; it was also sinking, as shown by getting out of focus. The seedlings were again placed in darkness for another hour, and the apex now crossed two divisions in 6 m. 18 s.; that is, at very nearly the same rate as before. After another interval of an hour in darkness, it crossed two divisions in 4 m. 15 s., therefore at a quicker rate. In the afternoon, after a longer interval in the dark, the apex was motionless, but after a time it recommenced moving, though slowly; perhaps the room was too cold. Judging from previous cases, there can hardly be a doubt that this seedling was circumnutating.
Solanum lycopersicum (Solaneae).—The movements of the hypocotyls of two seedling tomatoes were observed during seven hours, and there could be no doubt that both circumnutated. They were illuminated from above, but by an accident a little light entered on one side, and in the accompanying figure (Fig. 37) it may be seen that the hypocotyl moved to this side (the upper one in the figure), making small loops and zigzagging in its course. The movements of the cotyledons were also traced both on vertical and horizontal glasses; their angles with the horizon were likewise measured at various hours. They fell from 8.30 A.M. (October 17th) to about noon; then moved laterally in a zigzag line, and at about 4 P.M. began to rise; they continued to do so until 10.30 P.M., by which hour they stood vertically and were asleep. At what hour of the night or early morning they began to fall was not ascertained. Owing to the lateral movement shortly after mid-day, the descending and ascending lines did not coincide, and irregular ellipses were described during each 24 h. The regular periodicity of these movements is destroyed, as we shall hereafter see, if the seedlings are kept in the dark.
Fig. 37. Solanum lycopersicum: circumnutation of hypocotyl, with filament fixed across its summit, traced on horizontal glass, from 10 A.M. to 5 P.M. Oct. 24th. Illuminated obliquely from above. Movement of bead magnified about 35 times, here reduced to one-third of original scale.
Solanum palinacanthum.—Several arched hypocotyls rising nearly.2 of an inch above the ground, but with the cotyledons still buried beneath the surface, were observed, and the tracings showed that they circumnutated. Moreover, in several cases little open circular spaces