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 4 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 35o 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 [page 41] 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 [page 42] 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 14 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 60o, 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) (Cucurbitaceae).—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 53o, and at 10 P.M. they were in close contact, so that each had risen 26 1/2o. At noon, on the 29th, they included an angle of 118o, and at 10 P.M. an angle of 54o, so each had risen 32o. 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, [page 43] open more and more on each succeeding day, and rise each night about 30o; consequently during the first two nights of their life they stand vertically and come into contact.

Fig. 30. Lagenaria vulgaris: circumnutation of a cotyledon, 1 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 17 1/2o to 18o C. (63o to 65o 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 (Cucurbitaceae).—Two seedlings had opened [page 44] their cotyledons for the first time during the day,—one to the extent of 90o 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 90o 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 (Cacteae).—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 10o 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 [page 45] 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 (Compositae).—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 73o 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 [page 46] 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* 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 24 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. 23-24o 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

* 'Bot. Zeitung,' 1874, p. 837. [page 47]

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.

Ipomoea 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.* 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 3 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

* 'Movements and Habits of Climbing Plants,' p. 33, 1875. [page 48]

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 down in the course of 16 h. 20 m. (i.e. between 6.10 A.M. and 10.30 P.M.) thirteen times.

Fig 35. Ipomoea 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. 20o 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 30o and 35o; but three stood at about 50o and one at even 70o beneath the horizon. The blades of all these cotyledons had attained almost their full size, viz. from 1 to 1 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 [page 49] 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 Ipomoea likewise sink downwards late in the evening.

Cerinthe major (Boragineae).—The circumnutation of the hypocotyl of a young seedling with the cotyledons hardly

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.

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. [page 50]

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.

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 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. [page 51]

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 or cracks in the argillaceous sand which surrounded the arched hypocotyls were visible, and these appeared to have been made by the hypocotyls having bent first to one and then to another side whilst growing upwards. In two instances the vertical arches were observed to move to a considerable distance backwards from the point where the cotyledons lay buried; this movement, which has been noticed in some other cases, and which seems to aid in extracting the cotyledons from the buried seed-coats, is due to the commencement of the straightening of the hypocotyl. In order to prevent this latter movement, the two legs of an arch, the

Fig. 38. Solanum palinacanthum: circumnutation of an arched hypocotyl, just emerging from the ground, with the two legs tied together, traced in darkness on a horizontal glass, from 9.20 A.M. Dec. 17th to 8.30 A.M. 19th. Movement of bead magnified 13 times; but the filament, which was affixed obliquely to the crown of the arch, was of unusual length.

summit of which was on a level with the surface of the soil, were tied together; the earth having been previously removed to a little depth all round. The movement of the arch during 47 hours under these unnatural circumstances is exhibited in the annexed figure.

The cotyledons of some seedlings in the hot-house were horizontal about noon on December 13th; and at 10 P.M. had risen to an angle of 27o above the horizon; at 7 A.M. on the following [page 52] morning, before it was light, they had risen to 59o above the horizon; in the afternoon of the same day they were found again horizontal.

Beta vulgaris (Chenopodeae).—The seedlings are excessively sensitive to light, so that although on the first day they were uncovered only during two or three minutes at each observation, they all moved steadily towards the side of the room whence the light proceeded, and the tracings consisted only of slightly zigzag lines directed towards the light. On the next day the plants were placed in a completely darkened room, and at each observation were illuminated as much as possible from vertically above by a small wax taper. The annexed figure (Fig. 39) shows the movement of the hypocotyl during 9 h. under these circumstances. A second seedling was similarly observed at the same time, and the tracing had the same peculiar character, due to the hypocotyl often moving and returning in nearly parallel lines. The movement of a third hypocotyl differed greatly.

Fig. 39. Beta vulgaris: circumnutation of hypocotyl, with filament fixed obliquely across its summit, traced in darkness on horizontal glass, from 8.25 A.M. to 5.30 P.M. Nov. 4th. Movement of bead magnified 23 times, here reduced to one-third of original scale.

We endeavoured to trace the movements of the cotyledons, and for this purpose some seedlings were kept in the dark, but they moved in an abnormal manner; they continued rising from 8.45 A.M. to 2 P.M., then moved laterally, and from 3 to 6 P.M. descended; whereas cotyledons which have been exposed all the day to the light rise in the evening so as to stand vertically at night; but this statement applies only to young seedlings. For instance, six seedlings in the greenhouse had their cotyledons partially open for the first time on the morning of November 15th, and at 8.45 P.M. all were completely closed, so that they might properly be said to be asleep. Again, on the morning of November 27th, the cotyledons of four other seedlings, which were surrounded by a collar of brown paper so that they received light only from above, were open to the extent of 39o; at 10 P.M. they were completely closed; next morning (November 28th) at 6.45 A.M. whilst it was still dark, two of them [page 53] were partially open and all opened in the course of the morning; but at 10.20 P.M. all four (not to mention nine others which had been open in the morning and six others on another occasion) were again completely closed. On the morning of the 29th they were open, but at night only one of the four was closed, and this only partially; the three others had their cotyledons much more raised than during the day. On the night of the 30th the cotyledons of the four were only slightly raised.

Ricinus Borboniensis (Euphorbiaceae).—Seeds were purchased under the above name—probably a variety of the common castor-oil plant. As soon as an arched hypocotyl had risen clear above the ground, a filament was attached to the upper leg bearing the cotyledons which were still buried beneath the surface, and the movement of the bead was traced on a horizontal glass during a period of 34 h. The lines traced were strongly zigzag, and as the bead twice returned nearly parallel to its former course in two different directions, there could be no doubt that the arched hypocotyl circumnutated. At the close of the 34 h. the upper part began to rise and straighten itself, dragging the cotyledons out of the ground, so that the movements of the bead could no longer be traced on the glass.

Quercus (American sp.) (Cupuliferae).—Acorns of an American oak which had germinated at Kew were planted in a pot in the greenhouse. This transplantation checked their growth; but after a time one grew to a height of five inches, measured to the tips of the small partially unfolded leaves on the summit, and now looked vigorous. It consisted of six very thin internodes of unequal lengths. Considering these circumstances and the nature of the plant, we hardly expected that it would circumnutate; but the annexed figure (Fig. 40) shows that it did so in a conspicuous manner, changing its course many times and travelling in all directions during the 48 h. of observation. The figure seems to represent 5 or 6 irregular ovals or ellipses. The actual amount of movement from side to side (excluding one great bend to the left) was about .2 of an inch; but this was difficult to estimate, as owing to the rapid growth of the stem, the attached filament was much further from the mark beneath at the close than at the commencement of the observations. It deserves notice that the pot was placed in a north-east room within a deep box, the top of which was not at first covered up, so that the inside facing [page 54] the windows was a little more illuminated than the opposite side; and during the first morning the stem travelled to a greater distance in this direction (to the left in the figure) than it did afterwards when the box was completely protected from light.

Fig. 40. Quercus (American sp.): circumnutation of young stem, traced on horizontal glass, from 12.50 P.M. Feb. 22nd to 12.50 P.M. 24th. Movement of bead greatly magnified at first, but slightly towards the close of the observations—about 10 times on an average.

Quercus robur.—Observations were made only on the movements of the radicles from germinating acorns, which were allowed to grow downwards in the manner previously described, over plates of smoked glass, inclined at angles between 65o and 69o to the horizon. In four cases the tracks left were almost straight, but the tips had pressed sometimes with more and sometimes with less force on the glass, as shown by the varying thickness of the tracks and by little bridges of soot left across them. In the fifth case the track was slightly serpentine, that is, the tip had moved a little from side to side. In the sixth case (Fig. 41, A) it was plainly serpentine, and the tip had pressed almost equably on the glass in its whole course. In the seventh case (B) the tip had moved both laterally and had pressed [page 55] alternately with unequal force on the glass; so that it had moved a little in two planes at right angles to one another. In the eighth and last case (C) it had moved very little laterally, but had alternately left the glass and come into contact with it again. There can be no doubt that in the last four cases the radicle of the oak circumnutated whilst growing downwards.

Fig. 41. Quercus robur: tracks left on inclined smoked glass-plates by tips of radicles in growing downwards. Plates A and C inclined at 65o and plate B at 68o to the horizon.

Corylus avellana (Corylaceae).—The epicotyl breaks through the ground in an arched form; but in the specimen which was first examined, the apex had become decayed, and the epicotyl grew to some distance through the soil, in a tortuous, almost horizontal direction, like a root. In consequence of this injury it had emitted near the hypogean cotyledons two secondary shoots, and it was remarkable that both of these were arched, like the normal epicotyl in ordinary cases. The soil was removed from around one of these arched secondary shoots, and a glass filament was affixed to the basal leg. The whole was kept damp beneath a metal-box with a glass lid, and was thus illuminated only from above. Owing apparently to the lateral pressure of the earth being removed, the terminal and bowed-down part of the shoot began at once to move upwards, so that after 24 h. it formed a right angle with the lower part. This lower part, to which the filament was attached, also straightened itself, and moved a little backwards from the upper part. Consequently a long line was traced on the horizontal glass; and [page 56] this was in parts straight and in parts decidedly zigzag, indicating circumnutation.

On the following day the other secondary shoot was observed; it was a little more advanced in age, for the upper part, instead of depending vertically downwards, stood at an angle of 45o above the horizon. The tip of the shoot projected obliquely .4 of an inch above the ground, but by the close of our observations, which lasted 47 h., it had grown, chiefly towards its base, to a height of .85 of an inch. The filament was fixed transversely to the basal and almost upright half of the shoot, close beneath the lowest scale-like appendage. The circumnutating course pursued is shown in the accompanying figure (Fig. 42). The actual distance traversed from side to side was about .04 of an inch.

Fig. 42. Corylus avellana: circumnutation of a young shoot emitted from the epicotyl, the apex of which had been injured, traced on a horizontal glass, from 9 A.M. Feb. 2nd to 8 A.M. 4th. Movement of bead magnified about 27 times.

Pinus pinaster (Coniferae).—A young hypocotyl, with the tips of the cotyledons still enclosed within the seed-coats, was at first only .35 of an inch in height; but the upper part grew so rapidly that at the end of our observations it was .6 in height,

Fig. 43. Pinus pinaster: circumnutation of hypocotyl, with filament fixed across its summit, traced on horizontal glass, from 10 A.M. March 21st to 9 A.M. 23rd. Seedling kept in darkness. Movement of bead magnified about 35 times. [page 57]

and by this time the filament was attached some way down the little stem. From some unknown cause, the hypocotyl moved far towards the left, but there could be no doubt (Fig. 43) that it circumnutated. Another hypocotyl was similarly observed, and it likewise moved in a strongly zigzag line to the same side. This lateral movement was not caused by the attachment of the glass filaments, nor by the action of light; for no light was allowed to enter when each observation was made, except from vertically above.

The hypocotyl of a seedling was secured to a little stick; it bore nine in appearance distinct cotyledons, arranged in a circle. The movements of two nearly opposite ones were observed. The tip of one was painted white, with a mark placed below, and the figure described (Fig. 44, A) shows that it made an irregular

Fig. 44. Pinus pinaster: circumnutation of two opposite cotyledons, traced on horizontal glass in darkness, from 8.45 A.M. to 8.35 P.M. Nov. 25th. Movement of tip in A magnified about 22 times, here reduced to one-half of original scale.

circle in the course of about 8 h. during the night it travelled to a considerable distance in the direction indicated by the broken line. A glass filament was attached longitudinally to the other cotyledon, and this nearly completed (Fig, 44, B) an irregular circular figure in about 12 hours. During the night it also moved to a considerable distance, in the direction indicated by the broken line. The cotyledons therefore circumnutate independently of the movement of the hypocotyl. Although they moved much during the night, they did not approach each other so as to stand more vertically than during the day. [page 58]

Cycas pectinata (Cycadeae).—The large seeds of this plant in germinating first protrude a single leaf, which breaks through the ground with the petiole bowed into an arch and with the leaflets involuted. A leaf in this condition, which at the close of our observations was 2 inches in height, had its movements traced in a warm greenhouse by means of a glass filament bearing paper triangles attached across its tip. The tracing (Fig. 45) shows how large, complex, and rapid were the circum-

Fig. 45. Cycas pectinata: circumnutation of young leaf whilst emerging from the ground, feebly illuminated from above, traced on vertical glass, from 5 P.M. May 28th to 11 A.M. 31st. Movement magnified 7 times, here reduced to two-thirds of original scale.

nutating movements. The extreme distance from side to side which it passed over amounted to between .6 and .7 of an inch.

Canna Warscewiczii (Cannaceae).—A seedling with the plumule projecting one inch above the ground was observed, but not under fair conditions, as it was brought out of the hot-house and kept in a room not sufficiently warm. Nevertheless the tracing (Fig. 46) shows that it made two or three incomplete irregular circles or ellipses in the course of 48 hours. The plumule is straight; and this was the first instance observed [page 59] by us of the part that first breaks through the ground not being arched.

Fig. 46. Canna Warscewiczii: circumnutation of plumule with filament affixed obliquely to outer sheath-like leaf, traced in darkness on horizontal glass from 8.45 A.M. Nov. 9th to 8.10 A.M. 11th. Movement of bead magnified 6 times.

Allium cepa (Liliaceae).—The narrow green leaf, which protrudes from the seed of the common onion as a cotyledon,* breaks through the ground in the form of an arch, in the same manner as the hypocotyl or epicotyl of a dicotyledonous plant. Long after the arch has risen above the surface the apex remains within the seed-coats, evidently absorbing the still abundant contents. The summit or crown of the arch, when it first protrudes from the seed and is still buried beneath the ground, is simply rounded; but before it reaches the surface it is developed into a conical protuberance of a white colour (owing to the absence of chlorophyll), whilst the adjoining parts are green, with the epidermis apparently rather thicker and tougher than elsewhere. We may therefore conclude that this conical protuberance is a special adaptation for breaking through the ground,** and answers the same end as the knife-like white crest on the summit of the straight cotyledon of the Gramineae.

* This is the expression used by Sachs in his 'Text-book of Botany.'

** Haberlandt has briefly described ('Die Schutzeinrichtungen...Keimpflanze,' 1877, p. 77) this curious structure and the purpose which it subserves. He states that good figures of the cotyledon of the onion have been given by Tittmann and by Sachs in his 'Experimental Physiologie,' p. 93. [page 60]

After a time the apex is drawn out of the empty seed-coats, and rises up, forming a right angle, or more commonly a still larger angle with the lower part, and occasionally the whole becomes nearly straight. The conical protuberance, which originally formed the crown of the arch, is now seated on one side, and appears like a joint or knee, which from acquiring chlorophyll becomes green, and increases in size. In rarely or never becoming perfectly straight, these cotyledons differ remarkably from the ultimate condition of the arched hypocotyls or epicotyls of dicotyledons. It is, also, a singular circumstance that the attenuated extremity of the upper bent portion invariably withers and dies.

A filament, 1.7 inch in length, was affixed nearly upright beneath the knee to the basal and vertical portion of a cotyledon; and its movements were traced during 14 h. in the usual manner. The tracing here given (Fig. 47) indicates circumnutation. The movement of the upper part above the knee of the same cotyledon, which projected at about an angle of 45o above the horizon, was observed at the same time. A filament was not affixed to it, but a mark was placed beneath the apex, which was almost white from beginning to wither, and its movements were thus traced. The figure described resembled pretty closely that above given; and this shows that the chief seat of movement is in the lower or basal part of the cotyledon.

Fig. 47. Allium cepa: circumnutation of basal half of arched cotyledon, traced in darkness on horizontal glass, from 8.15 A.M. to 10 P.M. Oct. 31st. Movement of bead magnified about 17 times.

Asparagus officinalis (Asparageae).—The tip of a straight plumule or cotyledon (for we do not know which it should be called) was found at a depth of .1 inch beneath the surface, and the earth was then removed all round to the dept of .3 inch. a glass filament was affixed obliquely to it, and the movement of the bead, magnified 17 times, was traced in darkness. During the first 1 h. 15 m. the plumule moved to the right, and during the next two hours it returned in a roughly parallel but strongly zigzag course. From some unknown cause it had grown up through the soil in an inclined direction, and now through apogeotropism it moved during nearly 24 h. in [page 61] the same general direction, but in a slightly zigzag manner, until it became upright. On the following morning it changed its course completely. There can therefore hardly be a doubt that the plumule circumnutates, whilst buried beneath the ground, as much as the pressure of the surrounding earth will permit. The surface of the soil in the pot was now covered with a thin layer of very fine argillaceous sand, which was kept damp; and after the tapering seedlings had grown a few tenths of an inch in height, each was found surrounded by a little open space or circular crack; and this could be accounted for only by their having circumnutated and thus pushed away the sand on all sides; for there was no vestige of a crack in any other part.

In order to prove that there was circumnutation, the move-

Fig. 48. Asparagus officinalis: circumnutation of plumules with tips whitened and marks placed beneath, traced on a horizontal glass. A, young plumule; movement traced from 8.30 A.M. Nov. 30th to 7.15 A.M. next morning; magnified about 35 times. B, older plumule; movement traced from 10.15 A.M. to 8.10 P.M. Nov. 29th; magnified 9 times, but here reduced to one-half of original scale.

ments of five seedlings, varying in height from .3 inch to 2 inches, were traced. They were placed within a box and illuminated from above; but in all five cases the longer axes of the figures described were directed to nearly the same point; so that more light seemed to have come through the glass roof of the greenhouse on one side than on any other. All five tracings resembled each other to a certain extent, and it will suffice to give two of them. In A (Fig. 48) the seedling was only .45 of an [page 62] inch in height, and consisted of a single internode bearing a bud on its summit. The apex described between 8.30 A.M. and 10.20 P.M. (i.e. during nearly 14 hours) a figure which would probably have consisted of 3 ellipses, had not the stem been drawn to one side until 1 P.M., after which hour it moved backwards. On the following morning it was not far distant from the point whence it had first started. The actual amount of movement of the apex from side to side was very small, viz. about 1/18th of an inch. The seedling of which the movements are shown in Fig. 48, B, was 1 3/4 inch in height, and consisted of three internodes besides the bud on the summit. The figure, which was described during 10 h., apparently represents two irregular and unequal ellipses or circles. The actual amount of movement of the apex, in the line not influenced by the light, was .11 of an inch, and in that thus influenced .37 of an inch. With a seedling 2 inches in height it was obvious, even without the aid of any tracing, that the uppermost part of the stem bent successively to all points of the compass, like the stem of a twining plant. A little increase in the power of circumnutating and in the flexibility of the stem, would convert the common asparagus into a twining plant, as has occurred with one species in this genus, namely, A. scandens.

Phalaris Canariensis (Gramineae).—With the Gramineae the part which first rises above the ground has been called by some authors the pileole; and various views have been expressed on its homological nature. It is considered by some great authorities to be a cotyledon, which term we will use without venturing to express any opinion on the subject.* It consists in the present case of a slightly flattened reddish sheath, terminating upwards in a sharp white edge; it encloses a true green leaf, which protrudes from the sheath through a slit-like orifice, close beneath and at right angles to the sharp edge on the summit. The sheath is not arched when it breaks through the ground.

The movements of three rather old seedlings, about 1 inch in height, shortly before the protrusion of the leaves, were first traced. They were illuminated exclusively from above; for, as will hereafter be shown, they are excessively sensitive to the * We are indebted to the Rev. G. Henslow for an abstract of the views which have been held on this subject, together with references. [page 63]

action of light; and if any enters even temporarily on one side, they merely bend to this side in slightly zigzag lines. Of the three tracings one alone (Fig. 49) is here given. Had the observations been more frequent during the 12 h. two oval figures would have been described with their longer axes at right angles to one another. The actual amount of movement of the apex from side to side was about .3 of an inch. The figures described by the other two seedlings resembled to a certain extent the one here given.

Fig. 49. Phalaris Canariensis: circumnutation of a cotyledon, with a mark placed below the apex, traced on a horizontal glass, from 8.35 A.M. Nov. 26th to 8.45 A.M. 27th. Movement of apex magnified 7 times, here reduced to one-half scale.

A seedling which had just broken through the ground and projected only 1/20th of an inch above the surface, was next observed in the same manner as before. It was necessary to clear away the earth all round the seedling to a little depth in order to place a mark beneath the apex. The figure (Fig. 50) shows that the apex moved to one side, but changed its course ten times in the course of the ten hours of observation; so that there can be no doubt about its circumnutation. The cause of the general movement in one direction could hardly be attributed to the entrance of lateral light, as this was carefully guarded against; and we suppose it was in some manner connected with the removal of the earth round the little seedling.

Fig. 50. Phalaris Canariensis: circumnutation of a very young cotyledon, with a mark placed below the apex, traced on a horizontal glass, from 11.37 A.M. to 9.30 P.M. Dec. 13th. Movement of apex greatly magnified, here reduced to one-fourth of original scale.

Lastly, the soil in the same pot was searched with the aid of a lens, and the white knife-like apex of a seedling was found on an exact level with that of the surrounding surface. The soil was removed all round the apex to the depth of a quarter of an inch, the seed itself remaining covered. The pot, protected from lateral light, was placed under the micro- [page 64] scope with a micrometer eye-piece, so arranged that each division equalled 1/500th of an inch. After an interval of 30 m. the apex was observed, and it was seen to cross a little obliquely two divisions of the micrometer in 9 m. 15 s.; and after a few minutes it crossed the same space in 8 m. 50s. The seedling was again observed after an interval of three-quarters of an hour, and now the apex crossed rather obliquely two divisions in 10 m. We may therefore conclude that it was travelling at about the rate of 1/50th of an inch in 45 minutes. We may also conclude from these and the previous observations, that the seedlings of Phalaris in breaking through the surface of the soil circumnutate as much as the surrounding pressure will permit. This fact accounts (as in the case before given of the asparagus) for a circular, narrow, open space or crack being distinctly visible round several seedlings which had risen through very fine argillaceous sand, kept uniformly damp.

Fig. 51. Zea mays: circumnutation of cotyledon, traced on horizontal glass, from 8.30 A.M. Feb. 4th to 8 A.M. 6th. Movement of bead magnified on an average about 25 times.

Zea mays (Gramineae).—A glass filament was fixed obliquely to the summit of a cotyledon, rising .2 of an inch above the ground; but by the third morning it had grown to exactly thrice this height, so that the distance of the bead from the mark below was greatly increased, consequently the tracing (Fig. 51) was much more magnified on the first than on the second day. The upper part of the cotyledon changed its course by at least as much as a rectangle six times on each of the two days. The plant was illuminated by an obscure light from vertically above. This was a necessary precaution, as on the previous day we had traced the movements of cotyledons placed in a deep box, the inner side of which was feebly illuminated on one side from a distant north-east window, and at each observation by a wax taper held for a minute or two on the same side; and the result was that the cotyledons travelled all day long to this side, though making in their course some conspicuous flexures, from which fact alone we might have [page 65] concluded that they were circumnutating; but we thought it advisable to make the tracing above given.

Radicles.—Glass filaments were fixed to two short radicles, placed so as to stand almost upright, and whilst bending downwards through geotropism their courses were strongly zigzag; from this latter circumstance circumnutation might have been inferred, had not their tips become slightly withered after the first 24 h., though they were watered and the air kept very damp. Nine radicles were next arranged in the manner formerly described, so that in growing downwards they left tracks on smoked glass-plates, inclined at various angles between 45o and 80o beneath the horizon. Almost every one of these tracks offered evidence in their greater or less breadth in different parts, or in little bridges of soot being left, that the apex had come alternately into more and less close contact with the glass. In the accompanying figure (Fig. 52) we have an accurate copy of one such track. In two instances alone (and in these the plates were highly inclined) there was some evidence of slight lateral movement. We presume therefore that the friction of the apex on the smoked surface, little as this could have been, sufficed to check the movement from side to side of these delicate radicles.

Fig. 52. Zea mays: track left on inclined smoked glass-plate by tip of radicle in growing downwards.

Avena sativa (Gramineae).—A cotyledon, 1 inch in height, was placed in front of a north-east window, and the movement of the apex was traced on a horizontal glass during two days. It moved towards the light in a slightly zigzag line from 9 to 11.30 A.M. on October 15th; it then moved a little backwards and zigzagged much until 5 P.M., after which hour, and curing the night, it continued to move towards the window. On the following morning the same movement was continued in a nearly straight line until 12.40 P.M., when the sky remained until 2.35 extraordinarily dark from thunder-clouds. During this interval of 1 h. 55 m., whilst the light was obscure, it was interesting to observe how circumnutation overcame heliotropism, for the apex, instead of continuing to move towards the window in a slightly zigzag line, reversed its course four times, making two small narrow ellipses. A diagram of this case will be given in the chapter on Heliotropism. [page 66]

A filament was next fixed to a cotyledon only 1/4 of an inch in height, which was illuminated exclusively from above, and as it was kept in a warm greenhouse, it grew rapidly; and now there could be no doubt about its circumnutation, for it described a figure of 8 as well as two small ellipses in 5 hours.

Nephrodium molle (Filices).—A seedling fern of this species came up by chance in a flowerpot near its parent. The frond, as yet only slightly lobed, was only .16 of an inch in length and .2 in breadth, and was supported on a rachis as fine as a hair and .23 of an inch in height. A very thin glass filament, which projected for a length of .36 of an inch, was fixed to the end of the frond. The movement was so highly magnified that the figure (Fig. 53) cannot be fully trusted; but the frond was constantly moving in a complex manner, and the bead greatly changed its course eighteen times in the 12 hours of observation. Within half an hour it often returned in a line almost parallel to its former course. The greatest amount of movement occurred between 4 and 6 P.M. The circumnutation of this plant is interesting, because the species in the genus Lygodium are well known to circumnutate conspicuously and to twine round any neighbouring object.

Fig. 53. Nephrodium molle: circumnutation of very young frond, traced in darkness on horizontal glass, from 9 A.M. to 9 P.M. Oct. 30th. Movement of bead magnified 48 times.

Selaginella Kraussii (?) (Lycopodiaceae).—A very young plant, only .4 of an inch in height, had sprung up in a pot in the hot-house. An extremely fine glass filament was fixed to the end of the frond-like stem, and the movement of the bead traced on a horizontal glass. It changed its course several times, as shown in Fig. 54, whilst observed during 13 h. 15 m., and returned at night to a point not far distant from that whence it had started in the morning. There can be no doubt that this little plant circumnutated.

Fig. 54. Selaginella Kraussii (?): circumnutation of young plant, kept in darkness, traced from 8.45 A.M. to 10 P.M. Oct. 31st. [page 67]



CHAPTER II.

GENERAL CONSIDERATIONS ON THE MOVEMENTS AND GROWTH OF SEEDLING PLANTS.

Generality of the circumnutating movement—Radicles, their circumnutation of service—Manner in which they penetrate the ground—Manner in which hypocotyls and other organs break through the ground by being arched— Singular manner of germination in Megarrhiza, etc.—Abortion of cotyledons- -Circumnutation of hypocotyls and epicotyls whilst still buried and arched- -Their power of straightening themselves—Bursting of the seed-coats— Inherited effect of the arching process in hypogean hypocotyls— Circumnutation of hypocotyls and epicotyls when erect—Circumnutation of cotyledons—Pulvini or joints of cotyledons, duration of their activity, rudimentary in Oxalis corniculata, their development—Sensitiveness of cotyledons to light and consequent disturbance of their periodic movements- -Sensitiveness of cotyledons to contact.

THE circumnutating movements of the several parts or organs of a considerable number of seedling plants have been described in the last chapter. A list is here appended of the Families, Cohorts, Sub-classes, etc., to which they belong, arranged and numbered according to the classification adopted by Hooker.* Any one who will consider this list will see that the young plants selected for observation, fairly represent the whole vegetable series excepting the lowest cryptogams, and the movements of some of the latter when mature will hereafter be described. As all the seedlings which were observed, including Conifers, Cycads and Ferns, which belong to the most ancient

* As given in the 'General System of Botany,' by Le Maout and Decaisne, 1873. [page 68]

types amongst plants, were continually circumnutating, we may infer that this kind of movement is common to every seedling species.

SUB-KINGDOM I.—Phaenogamous Plants.

Class I.—DICOTYLEDONS.

Sub-class I.—Angiosperms. Family. Cohort. 14. Cruciferae. II. PARIETALES. 26. Caryophylleae. IV. CARYOPHYLLALES. 36. Malvaceae. VI MALVALES. 41. Oxalideae. VII. GERANIALES. 49. Tropaeoleae. DITTO 52. Aurantiaceae. DITTO 70. Hippocastaneae. X. SAPINDALES. 75. Leguminosae. XI. ROSALES. 106. Cucurbitaceae. XII. PASSIFLORALES. 109. Cacteae. XIV. FICOIDALES. 122. Compositae. XVII. ASTRALES. 135. Primulaceae. XX. PRIMULALES. 145. Asclepiadeae. XXII. GENTIANALES. 151. Convolvulaceae. XXIII. POLEMONIALES. 154. Boragineae. DITTO 156. Nolaneae. DITTO 157. Solaneae. XXIV. SOLANALES. 181. Chenopodieae. XXVII. CHENOPODIALES. 202. Euphorbiaceae. XXXII. EUPHORBIALES. 211. Cupuliferae. XXXVI. QUERNALES. 212. Corylaceae. DITTO

Sub-class II.—Gymnosperms. 223. Coniferae. 224. Cycadeae.

Class II.—MONOCOTYLEDONS. 2. Cannaceae. II. AMOMALES. 34. Liliaceae. XI. LILIALES. 41. Asparageae. DITTO 55. Gramineae. XV. GLUMALES.

SUB-KINGDOM II.—Cryptogamic Plants.

1. Filices. I. FILICALES. 6. Lycopodiaceae. DITTO [page 69]

Radicles.—In all the germinating seeds observed by us, the first change is the protrusion of the radicle, which immediately bends downwards and endeavours to penetrate the ground. In order to effect this, it is almost necessary that the seed should be pressed down so as to offer some resistance, unless indeed the soil is extremely loose; for otherwise the seed is lifted up, instead of the radicle penetrating the surface. But seeds often get covered by earth thrown up by burrowing quadrupeds or scratching birds, by the castings of earth-worms, by heaps of excrement, the decaying branches of trees, etc., and will thus be pressed down; and they must often fall into cracks when the ground is dry, or into holes. Even with seeds lying on the bare surface, the first developed root-hairs, by becoming attached to stones or other objects on the surface, are able to hold down the upper part of the radicle, whilst the tip penetrates the ground. Sachs has shown* how well and closely root-hairs adapt themselves by growth to the most irregular particles in the soil, and become firmly attached to them. This attachment seems to be effected by the softening or liquefaction of the outer surface of the wall of the hair and its subsequent consolidation, as will be on some future occasion more fully described. This intimate union plays an important part, according to Sachs, in the absorption of water and of the inorganic matter dissolved in it. The mechanical aid afforded by the root-hairs in penetrating the ground is probably only a secondary service.

The tip of the radicle, as soon as it protrudes from the seed-coats, begins to circumnutate, and the whole

* 'Physiologie Vgtale,' 1868, pp. 199, 205. [page 70]

growing part continues to do so, probably for as long as growth continues. This movement of the radicle has been described in Brassica, Aesculus, Phaseolus, Vicia, Cucurbita, Quercus and Zea. The probability of its occurrence was inferred by Sachs,* from radicles placed vertically upwards being acted on by geotropism (which we likewise found to be the case), for if they had remained absolutely perpendicular, the attraction of gravity could not have caused them to bend to any one side. Circumnutation was observed in the above specified cases, either by means of extremely fine filaments of glass affixed to the radicles in the manner previously described, or by their being allowed to grow downwards over inclined smoked glass-plates, on which they left their tracks. In the latter cases the serpentine course (see Figs. 19, 21, 27, 41) showed unequivocally that the apex had continually moved from side to side. This lateral movement was small in extent, being in the case of Phaseolus at most about 1 mm. from a medial line to both sides. But there was also movement in a vertical plane at right angles to the inclined glass-plates. This was shown by the tracks often being alternately a little broader and narrower, due to the radicles having alternately pressed with greater and less force on the plates. Occasionally little bridges of soot were left across the tracks, showing that the apex had at these spots been lifted up. This latter fact was especially apt to occur * 'Ueber das Wachsthum der Wurzeln: Arbeiten des bot. Instituts in Wrzburg,' Heft iii. 1873, p. 460. This memoir, besides its intrinsic and great interest, deserves to be studied as a model of careful investigation, and we shall have occasion to refer to it repeatedly. Dr. Frank had previously remarked ('Beitrge zur Pflanzenphysiologie, 1868, p. 81) on the fact of radicles placed vertically upwards being acted on by geotropism, and he explained it by the supposition that their growth was not equal on all sides.

[page 71] when the radicle instead of travelling straight down the glass made a semicircular bend; but Fig. 52 shows that this may occur when the track is rectilinear. The apex by thus rising, was in one instance able to surmount a bristle cemented across an inclined glass-plate; but slips of wood only 1/40 of an inch in thickness always caused the radicles to bend rectangularly to one side, so that the apex did not rise to this small height in opposition to geotropism.

In those cases in which radicles with attached filaments were placed so as to stand up almost vertically, they curved downwards through the action of geotropism, circumnutating at the same time, and their courses were consequently zigzag. Sometimes, however, they made great circular sweeps, the lines being likewise zigzag.

Radicles closely surrounded by earth, even when this is thoroughly soaked and softened, may perhaps be quite prevented from circumnutating. Yet we should remember that the circumnutating sheath-like cotyledons of Phalaris, the hypocotyls of Solanum, and the epicotyls of Asparagus formed round themselves little circular cracks or furrows in a superficial layer of damp argillaceous sand. They were also able, as well as the hypocotyls of Brassica, to form straight furrows in damp sand, whilst circumnutating and bending towards a lateral light. In a future chapter it will be shown that the rocking or circumnutating movement of the flower-heads of Trifolium subterraneum aids them in burying themselves. It is therefore probable that the circumnutation of the tip of the radicle aids it slightly in penetrating the ground; and it may be observed in several of the previously given diagrams, that the movement is more strongly pronounced in radicles when they first [page 72] protrude from the seed than at a rather later period; but whether this is an accidental or an adaptive coincidence we do not pretend to decide. Nevertheless, when young radicles of Phaseolus multiflorus were fixed vertically close over damp sand, in the expectation that as soon as they reached it they would form circular furrows, this did not occur,—a fact which may be accounted for, as we believe, by the furrow being filled up as soon as formed by the rapid increase of thickness in the apex of the radicle. Whether or not a radicle, when surrounded by softened earth, is aided in forming a passage for itself by circumnutating, this movement can hardly fail to be of high importance, by guiding the radicle along a line of least resistance, as will be seen in the next chapter when we treat of the sensibility of the tip to contact. If, however, a radicle in its downward growth breaks obliquely into any crevice, or a hole left by a decayed root, or one made by the larva of an insect, and more especially by worms, the circumnutating movement of the tip will materially aid it in following such open passage; and we have observed that roots commonly run down the old burrows of worms.*

When a radicle is placed in a horizontal or inclined position, the terminal growing part, as is well known, bends down towards the centre of the earth; and Sachs* has shown that whilst thus bending, the growth of the lower surface is greatly retarded, whilst that

* See, also, Prof. Hensen's statements ('Zeitschrift fr Wissen, Zool.,' B. xxviii. p. 354, 1877) to the same effect. He goes so far as to believe that roots are able to penetrate the ground to a great depth only by means of the burrows made by worms.

* 'Arbeiten des bot. Inst. Wrzburg,' vol. i. 1873, p. 461. See also p. 397 for the length of the growing part, and p. 451 on the force of geotropism. [page 73]

of the upper surface continues at the normal rate, or may be even somewhat increased. He has further shown by attaching a thread, running over a pulley, to a horizontal radicle of large size, namely that of the common bean, that it was able to pull up a weight of only one gramme, or 15.4 grains. We may therefore conclude that geotropism does not give a radicle force sufficient to penetrate the ground, but merely tells it (if such an expression may be used) which course to pursue. Before we knew of Sachs' more precise observations we covered a flat surface of damp sand with the thinnest tin-foil which we could procure (.02 to .03 mm., or .00012 to .00079 of an inch in thickness), and placed a radicle close above, in such a position that it grew almost perpendicularly downwards. When the apex came into contact with the polished level surface it turned at right angles and glided over it without leaving any impression; yet the tin-foil was so flexible, that a little stick of soft wood, pointed to the same degree as the end of the radicle and gently loaded with a weight of only a quarter of an ounce (120 grains) plainly indented the tin-foil.

Radicles are able to penetrate the ground by the force due to their longitudinal and transverse growth; the seeds themselves being held down by the weight of the superincumbent soil. In the case of the bean the apex, protected by the root-cap, is sharp, and the growing part, from 8 to 10 mm. in length, is much more rigid, as Sachs has proved, than the part immediately above, which has ceased to increase in length. We endeavoured to ascertain the downward pressure of the growing part, by placing germinating beans between two small metal plates, the upper one of which was loaded with a known weight; and the [page 74] radicle was then allowed to grow into a narrow hole in wood, 2 or 3 tenths of an inch in depth, and closed at the bottom. The wood was so cut that the short space of radicle between the mouth of the hole and the bean could not bend laterally on three sides; but it was impossible to protect the fourth side, close to the bean. Consequently, as long as the radicle continued to increase in length and remained straight, the weighted bean would be lifted up after the tip had reached the bottom of the shallow hole. Beans thus arranged, surrounded by damp sand, lifted up a quarter of a pound in 24 h. after the tip of the radicle had entered the hole. With a greater weight the radicles themselves always became bent on the one unguarded side; but this probably would not have occurred if they had been closely surrounded on all sides by compact earth. There was, however, a possible, but not probable, source of error in these trials, for it was not ascertained whether the beans themselves go on swelling for several days after they have germinated, and after having been treated in the manner in which ours had been; namely, being first left for 24 h. in water, then allowed to germinate in very damp air, afterwards placed over the hole and almost surrounded by damp sand in a closed box.

Fig. 55. Outline of piece of stick (reduced to one-half natural size) with a hole through which the radicle of a bean grew. Thickness of stick at narrow end .08 inch, at broad end .16; depth of hole .1 inch. We succeeded better in ascertaining the force exerted transversely by these radicles. Two were so placed as to penetrate small holes made in little sticks, one of which was cut into the shape here exactly copied (Fig. 55). The short end of the stick beyond the hole was purposely split, but not the opposite [page 75] end. As the wood was highly elastic, the split or fissure closed immediately after being made. After six days the stick and bean were dug out of the damp sand, and the radicle was found to be much enlarged above and beneath the hole. The fissure which was at first quite closed, was now open to a width of 4 mm.; as soon as the radicle was extracted, it immediately closed to a width of 2 mm. The stick was then suspended horizontally by a fine wire passing through the hole lately filled by the radicle, and a little saucer was suspended beneath to receive the weights; and it required 8 lbs. 8 ozs. to open the fissure to the width of 4 mm.— that is, the width before the root was extracted. But the part of the radicle (only .1 of an inch in length) which was embedded in the hole, probably exerted a greater transverse strain even than 8 lbs. 8 ozs., for it had split the solid wood for a length of rather more than a quarter of an inch (exactly .275 inch), and this fissure is shown in Fig. 55. A second stick was tried in the same manner with almost exactly the same result.

Fig. 56. Wooden pincers, kept closed by a spiral brass spring, with a hole (.14 inch in diameter and .6 inch in depth) bored through the narrow closed part, through which a radicle of a bean was allowed to grow. Temp. 50o - 60o F.

We then followed a better plan. Holes were bored near the narrow end of two wooden clips or pincers (Fig. 56), kept closed by brass spiral springs. Two radicles in damp sand were allowed to grow through these holes. The [page 76] pincers rested on glass-plates to lessen the friction from the sand. The holes were a little larger (viz..14 inch) and considerably deeper (viz..6 inch) than in the trials with the sticks; so that a greater length of a rather thicker radicle exerted a transverse strain. After 13 days they were taken up. The distance of two dots (see the figure) on the longer ends of the pincers was now carefully measured; the radicles were then extracted from the holes, and the pincers of course closed. They were then suspended horizontally in the same manner as were the bits of sticks, and a weight of 1500 grams (or 3 pounds 4 ounces) was necessary with one of the pincers to open them to the same extent as had been effected by the transverse growth of the radicle. As soon as this radicle had slightly opened the pincers, it had grown into a flattened form and had escaped a little beyond the hole; its diameter in one direction being 4.2 mm., and at rightangles 3.5 mm. If this escape and flattening could have been prevented, the radicle would probably have exerted a greater strain than the 3 pounds 4 ounces. With the other pincers the radicle escaped still further out of the hole; and the weight required to open them to the same extent as had been effected by the radicle, was only 600 grams.

With these facts before us, there seems little difficulty in understanding how a radicle penetrates the ground. The apex is pointed and is protected by the root-cap; the terminal growing part is rigid, and increases in length with a force equal, as far as our observations can be trusted, to the pressure of at least a quarter of a pound, probably with a much greater force when prevented from bending to any side by the surrounding earth. Whilst thus increasing in length it increases in thickness, pushing away the damp [page 77] earth on all sides, with a force of above 8 pounds in one case, of 3 pounds in another case. It was impossible to decide whether the actual apex exerts, relatively to its diameter, the same transverse strain as the parts a little higher up; but there seems no reason to doubt that this would be the case. The growing part therefore does not act like a nail when hammered into a board, but more like a wedge of wood, which whilst slowly driven into a crevice continually expands at the same time by the absorption of water; and a wedge thus acting will split even a mass of rock.

Manner in which Hypocotyls, Epicotyls, etc., rise up and break through the ground.—After the radicle has penetrated the ground and fixed the seed, the hypocotyls of all the dicotyledonous seedlings observed by us, which lift their cotyledons above the surface, break through the ground in the form of an arch. When the cotyledons are hypogean, that is, remain buried in the soil, the hypocotyl is hardly developed, and the epicotyl or plumule rises in like manner as an arch through the ground. In all, or at least in most of such cases, the downwardly bent apex remains for a time enclosed within the seed-coats. With Corylus avellena the cotyledons are hypogean, and the epicotyl is arched; but in the particular case described in the last chapter its apex had been injured, and it grew laterally through the soil like a root; and in consequence of this it had emitted two secondary shoots, which likewise broke through the ground as arches.

Cyclamen does not produce any distinct stem, and only a single cotyledon appears at first;* its petiole

* This is the conclusion arrived at by Dr. H. Gressner ('Bot. Zeitung,' 1874, p. 837), who maintains that what has been considered by other botanists as the first true leaf is really the second cotyledon, which is greatly delayed in its development. [page 78]

breaks through the ground as an arch (Fig. 57). Abronia has only a single fully developed cotyledon, but in this case it is the hypocotyl which first emerges and is arched. Abronia umbellata, however, presents this peculiarity, that the enfolded blade of the one developed cotyledon (with the enclosed endosperm) whilst still beneath the surface has its apex upturned and parallel to the descending leg of the arched hypocotyl; but it is dragged out of the ground by the continued growth of the hypocotyl, with the apex pointing downward. With Cycas pectinata the cotyledons are hypogean, and a true leaf first breaks through the ground with its petiole forming an arch.

Fig. 57. Cyclamen Persicum: seedling, figure enlarged: c, blade of cotyledon, not yet expanded, with arched petiole beginning to straighten itself; h, hypocotyl developed into a corm; r, secondary radicles.

Fig. 58. Acanthus mollis: seedling with the hypogean cotyledon on the near side removed and the radicles cut off; a, blade of first leaf beginning to expand, with petiole still partially arched; b, second and opposite leaf, as yet very imperfectly developed; c, hypogean cotyledon on the opposite side.

In the genus Acanthus the cotyledons are likewise hypogean. In A. mollis, a single leaf first breaks through the ground with its petiole arched, and with the opposite leaf much less developed, short, straight, of a yellowish colour, and with the petiole at first not half as thick as that of the other. The undeveloped leaf is protected by standing beneath its arched fellow; and it is an instruc- [page 79] tive fact that it is not arched, as it has not to force for itself a passage through the ground. In the accompanying sketch (Fig. 58) the petiole of the first leaf has already partially straightened itself, and the blade is beginning to unfold. The small second leaf ultimately grows to an equal size with the first, but this process is effected at very different rates in different individuals: in one instance the second leaf did not appear fully above the ground until six weeks after the first leaf. As the leaves in the whole family of the Acanthaceae stand either opposite one another or in whorls, and as these are of equal size, the great inequality between the first two leaves is a singular fact. We can see how this inequality of development and the arching of the petiole could have been gradually acquired, if they were beneficial to the seedlings by favouring their emergence; for with A. candelabrum, spinosus, and latifolius there was a great variability in the inequality between the two first leaves and in the arching of their petioles. In one seedling of A. candelabrum the first leaf was arched and nine times as long as the second, which latter consisted of a mere little, yellowish-white, straight, hairy style. In other seedlings the difference in length between the two leaves was as 3 to 2, or as 4 to 3, or as only .76 to .62 inch. In these latter cases the first and taller leaf was not properly arched. Lastly, in another seedling there was not the least difference in size between the two first leaves, and both of them had their petioles straight; their laminae were enfolded and pressed against each other, forming a lance or wedge, by which means they had broken through the ground. Therefore in different individuals of this same species of Acanthus the first pair of leaves breaks through the ground by two widely different methods; and if [page 80] either had proved decidedly advantageous or disadvantageous, one of them no doubt would soon have prevailed.

Asa Gray has described* the peculiar manner of germination of three widely different plants, in which the hypocotyl is hardly at all developed. These were therefore observed by us in relation to our present subject.

Delphinium nudicaule.—The elongated petioles of the two cotyledons are confluent (as are sometimes their blades at the base), and they break through the ground as an arch. They thus resemble in a most deceptive manner a hypocotyl. At first they are solid, but after a time become tubular; and the basal part beneath the ground is enlarged into a hollow chamber, within which the young leaves are developed without any prominent plumule. Externally root-hairs are formed on the confluent petioles, either a little above, or on a level with, the plumule. The first leaf at an early period of its growth and whilst within the chamber is quite straight, but the petiole soon becomes arched; and the swelling of this part (and probably of the blade) splits open one side of the chamber, and the leaf then emerges. The slit was found in one case to be 3.2 mm. in length, and it is seated on the line of confluence of the two petioles. The leaf when it first escapes from the chamber is buried beneath the ground, and now an upper part of the petiole near the blade becomes arched in the usual manner. The second leaf comes out of the slit either straight or somewhat arched, but afterwards the upper part of the petiole,—certainly in some, and we believe in all cases,—arches itself whilst forcing a passage through the soil.

* 'Botanical Text-Book,' 1879, p. 22. [page 81]

Megarrhiza Californica.—The cotyledons of this Gourd never free themselves from the seed-coats and are hypogean. Their petioles are completely confluent, forming a tube which terminates downwards in a little solid point, consisting of a minute radicle and hypocotyl, with the likewise minute plumule enclosed within the base of the tube. This structure was well exhibited in an abnormal specimen, in which one of the two cotyledons failed to produce a petiole, whilst the other produced one consisting of an open semicylinder ending in a sharp point, formed of the parts just described. As soon as the confluent petioles protrude from the seed they bend down, as they are strongly geotropic, and penetrate the ground. The seed itself retains its original position, either on the surface or buried at some depth, as the case may be. If, however, the point of the confluent petioles meets with some obstacle in the soil, as appears to have occurred with the seedlings described and figured by Asa Gray,* the cotyledons are lifted up above the ground. The petioles are clothed with root-hairs like those on a true radicle, and they likewise resemble radicles in becoming brown when immersed in a solution of permanganate of potassium. Our seeds were subjected to a high temperature, and in the course of three or four days the petioles penetrated the soil perpendicularly to a depth of from 2 to 2 inches; and not until then did the true radicle begin to grow. In one specimen which was closely observed, the petioles in 7 days after their first protrusion attained a length of 2 inches, and the radicle by this time had also become well developed. The plumule, still enclosed within the tube, was now

* 'American Journal of Science,' vol. xiv. 1877, p. 21. [page 82]

.3 inch in length, and was quite straight; but from having increased in thickness it had just begun to split open the lower part of the petioles on one side, along the line of their confluence. By the following morning the upper part of the plumule had arched itself into a right angle, and the convex side or elbow had thus been forced out through the slit. Here then the arching of the plumule plays the same part as in the case of the petioles of the Delphinium. As the plumule continued to grow, the tip became more arched, and in the course of six days it emerged through the 2 inches of superincumbent soil, still retaining its arched form. After reaching the surface it straightened itself in the usual manner. In the accompanying figure (Fig. 58, A) we have a sketch of a seedling in this advanced state of development; the surface of the ground being represented by the line G...........G.

Fig. 58, A. Megarrhiza Californica: sketch of seedling, copied from Asa Gray, reduced to one-half scale: c, cotyledons within seed-coats; p, the two confluent petioles; h and r, hypocotyl and radicle; p1, plumule; G..........G, surface of soil.

The germination of the seeds in their native Californian home proceeds in a rather different manner, as we infer from an interesting letter from Mr. Rattan, sent to us by Prof. Asa Gray. The petioles protrude from the seeds soon after the autumnal rains, and penetrate the ground, generally in a vertical direction, to a depth of from 4 to even 6 inches. they were found in this state by Mr. Rattan during the Christmas vacation, with the plu- [page 83] mules still enclosed within the tubes; and he remarks that if the plumules had been at once developed and had reached the surface (as occurred with our seeds which were exposed to a high temperature), they would surely have been killed by the frost. As it is, they lie dormant at some depth beneath the surface, and are thus protected from the cold; and the root-hairs on the petioles would supply them with sufficient moisture. We shall hereafter see that many seedlings are protected from frost, but by a widely different process, namely, by being drawn beneath the surface by the contraction of their radicles. We may, however, believe that the extraordinary manner of germination of Megarrhiza has another and secondary advantage. The radicle begins in a few weeks to enlarge into a little tuber, which then abounds with starch and is only slightly bitter. It would therefore be very liable to be devoured by animals, were it not protected by being buried whilst young and tender, at a depth of some inches beneath the surface. Ultimately it grows to a huge size.

Ipomoea leptophylla.—In most of the species of this genus the hypocotyl is well developed, and breaks through the ground as an arch. But the seeds of the present species in germinating behave like those of Megarrhiza, excepting that the elongated petioles of the cotyledons are not confluent. After they have protruded from the seed, they are united at their lower ends with the undeveloped hypocotyl and undeveloped radicle, which together form a point only about .1 inch in length. They are at first highly geotropic, and penetrate the ground to a depth of rather above half an inch. The radicle then begins to grow. On four occasions after the petioles had grown for a short distance vertically downwards, they [page 84] were placed in a horizontal position in damp air in the dark, and in the course of 4 hours they again became curved vertically downwards, having passed through 90o in this time. But their sensitiveness to geotropism lasts for only 2 or 3 days; and the terminal part alone, for a length of between .2 and .4 inch, is thus sensitive. Although the petioles of our specimens did not penetrate the ground to a greater depth than about inch, yet they continued for some time to grow rapidly, and finally attained the great length of about 3 inches. The upper part is apogeotropic, and therefore grows vertically upwards, excepting a short portion close to the blades, which at an early period bends downwards and becomes arched, and thus breaks through the ground. Afterwards this portion straightens itself, and the cotyledons then free themselves from the seed-coats. Thus we here have in different parts of the same organ widely different kinds of movement and of sensitiveness; for the basal part is geotropic, the upper part apogeotropic, and a portion near the blades temporarily and spontaneously arches itself. The plumule is not developed for some little time; and as it rises between the bases of the parallel and closely approximate petioles of the cotyledons, which in breaking through the ground have formed an almost open passage, it does not require to be arched and is consequently always straight. Whether the plumule remains buried and dormant for a time in its native country, and is thus protected from the cold of winter, we do not know. The radicle, like that of the Megarrhiza, grows into a tuber-like mass, which ultimately attains a great size. So it is with Ipomoea pandurata, the germination of which, as Asa Gray informs us, resembles that of I. leptophylla.

The following case is interesting in connection with [page 85] the root-like nature of the petioles. The radicle of a seedling was cut off, as it was completely decayed, and the two now separated cotyledons were planted. They emitted roots from their bases, and continued green and healthy for two months. The blades of both then withered, and on removing the earth the bases of the petioles (instead of the radicle) were found enlarged into little tubers. Whether these would have had the power of producing two independent plants in the following summer, we do not know.

In Quercus virens, according to Dr. Engelmann,* both the cotyledons and their petioles are confluent. The latter grow to a length "of an inch or even more;" and, if we understand rightly, penetrate the ground, so that they must be geotropic. The nutriment within the cotyledons is then quickly transferred to the hypocotyl or radicle, which thus becomes developed into a fusiform tuber. The fact of tubers being formed by the foregoing three widely distinct plants, makes us believe that their protection from animals at an early age and whilst tender, is one at least of the advantages gained by the remarkable elongation of the petioles of the cotyledons, together with their power of penetrating the ground like roots under the guidance of geotropism.

The following cases may be here given, as they bear on our present subject, though not relating to seedlings. The flower-stem of the parasitic Lathraea squamaria, which is destitute of true leaves, breaks through the ground as an arch;** so does the flower-

* 'Transact. St. Louis Acad. Science,' vol. iv. p. 190.

** The passage of the flower-stem of the Lathraea through the ground cannot fail to be greatly facilitated by the extraordinary quantity of water secreted at this period of the year by the subter- [[page 86]] ranean scale-like leaves; not that there is any reason to suppose that the secretion is a special adaptation for this purpose: it probably follows from the great quantity of sap absorbed in the early spring by the parasitic roots. After a long period without any rain, the earth had become light-coloured and very dry, but it was dark-coloured and damp, even in parts quite wet, for a distance of at least six inches all round each flower-stem. The water is secreted by glands (described by Cohn, 'Bericht. Bot. Sect. der Schlesischen Gesell.,' 1876, p. 113) which line the longitudinal channels running through each scale-like leaf. A large plant was dug up, washed so as to remove the earth, left for some time to drain, and then placed in the evening on a dry glass-plate, covered with a bell-glass, and by next morning it had secreted a large pool of water. The plate was wiped dry, and in the course of the succeeding 7 or 8 hours another little pool was secreted, and after 16 additional hours several large drops. A smaller plant was washed and placed in a large jar, which was left inclined for an hour, by which time no more water drained off. The jar was then placed upright and closed: after 23 hours two drachms of water were collected from the bottom, and a little more after 25 additional hours. The flower-stems were now cut off, for they do not secrete, and the subterranean part of the plant was found to weigh 106.8 grams (1611 grains), and the water secreted during the 48 hours weighed 11.9 grams (183 grains),—that is, one-ninth of the whole weight of the plant, excluding the flower-stems. We should remember that plants in a state of nature would probably secrete in 48 hours much more than the above large amount, for their roots would continue all the time absorbing sap from the plant on which they were parasitic. [page 86]

stem of the parasitic and leafless Monotropa hypopitys. With Helleborus niger, the flower-stems, which rise up independently of the leaves, likewise break through the ground as arches. This is also the case with the greatly elongated flower-stems, as well as with the petioles of Epimedium pinnatum. So it is with the petioles of Ranunculus ficaria, when they have to break through the ground, but when they arise from the summit of the bulb above ground, they are from the first quite straight; and this is a fact which deserves notice. The rachis of the bracken fern (Pteris aquilina), and of some, probably many, other ferns, likewise rises above ground under the form of an arch. No doubt other analogous instances could be found by careful search. In all ordinary cases of bulbs, rhizomes, [page 87] root-stocks, etc., buried beneath the ground, the surface is broken by a cone formed by the young imbricated leaves, the combined growth of which gives them force sufficient for the purpose.