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 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.
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.
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 27° above the horizon; at 7 A.M. on the following morning, before it was light, they had risen to 59° 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 39°; 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 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 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 65° and 69° 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 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 65° and plate B at 68° 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 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 45° 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 (Coniferæ).—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, 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.
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.
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 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.
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.
Cycas pectinata (Cycadeæ).—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 2½ 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 circumnutating movements. The extreme distance from side to side which it passed over amounted to between.6 and.7 of an inch.
Fig. 45. Cycas pectinata: circumnutation of young leaf whilst emerging from the ground, feebly illuminated from above, traced on vertical times, here reduced to two-thirds of original scale.
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 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,[4] 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,[5] and answers the same end as the knife-like white crest on the summit of the straight cotyledon of the Gramineæ.
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.
[4] This is the expression used by Sachs in his ‘Text-book of Botany.’
[5] Haberlandt has briefly described (‘Die 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.
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 45° 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 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 movements 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 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 3½ 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.
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.
Phalaris Canariensis (Gramineæ).—With the Gramineæ 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.[6] 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.
[6] We are indebted to the Rev. G. Henslow for an abstract of the views which have been held on this subject, together with references.
The movements of three rather old seedlings, about 1½ 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 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 microscope 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 (Gramineæ).—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 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 45° and 80° 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 (Gramineæ).—A cotyledon, 1½ 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.
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 5½ 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 (?) (Lycopodiaceæ).—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.
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.[1] 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 types amongst plants, were continually circumnutating, we may infer that this kind of movement is common to every seedling species.
[1] As given in the ‘General System of Botany,’ by Le Maout and Decaisne, 1873.
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. Cucurbitaceæ. XII. PASSIFLORALES. 109. Cacteæ. XIV.
FICOIDALES. 122. Compositæ. 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 Class II.—MONOCOTYLEDONS. 2. Cannaceae. II. AMOMALES. 34. Liliaceae.
XI. LILIALES. 41. Asparageae. DITTO 55. Gramineæ. XV. GLUMALES.
1. Filices. I. FILICALES. 6. Lycopodiaceæ. DITTO 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[2] 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 growing part continues to do so, probably for as long as growth continues. This movement of the radicle has been described in Brassica,