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traced on Oct 27th magnified about 56 40 times. This frond was originally inclined considerably upwards, having been placed before [insertion:] Of course the filament increased the movement, considerably, the movement of the frond end was greatly magnified a N. E window on Oct 19th at 9° 15' a.m it continued to sink till evening of Oct 24th, when filament fell off. (See tracing) As the frond kept on sinking day night probably not due to apheliotropism— either epinasty or geotropism.— As course was so
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monocotyledons p. 37 leaves of grow in darkness, probably some connection with the fact of them circumnutating in darkness.) [in margin:] Keep — Put in Chapter wh I speak of Darkness Light deciding curvature p. 30 31. germination of seeds in dark— radicles short. p. 32 shoots in darkness grow vertically this looks as if epinasty failed, for de Vries wd have discovered apheliotropism.— [Movement in plants, p. 443: It must be the result of some periodical change in the conditions to which they are subjected
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March 20' 1879— Pot with Cyclamen Persica Pot with long pod─ peduncles curved to ground — pot laid horizontally in few days peduncles all curved bent at right angles to former course— Possibly may be weight of Pod — or possibly Apheliotropism — [49
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All these cases indicate that guide to bending down is apheliotropism The circumnutation is going in Geotropic [illeg] (The fact of not bending down in dark cupboard shown not weight) the pods occupied— Pods were cut off 2 peduncle, these became a little bowed down but they tend to wither soon when pods cut off. — Another pot was placed horizontally in dark cupboard on March 20th on 26th none except perhaps one very slightly were turned down. On 26th brought back kept in same position into
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now turning up— if so hy epinasty Sept 17th 12° 30' uncovered pots of the 2 without thr threads to sticks / some of flowers on further ones are horizontal quite white to see whether apheliotropism can cause them to descend — no (over) [41v
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CUL-DAR209.7.133
Draft:
[1878].11.02--[1878].11.05
Tropaeolum majus / Draft of Cross and self fertilisation
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The Complete Work of Charles Darwin Online [133] Tropæolum majus— plants bow will lean to Light in greenhouse slew a little from them — reversed Pot placed before N.E. window — black on all other sides top. — filament with triangle fixed to penultimate internode— traced on Horizontal glass — Height of Plants? —Length of internodes? Distance from glass Left Hand [data not transcribed] Apheliotropism Right Hand [data not transcribed] (Used) [133v
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The Complete Work of Charles Darwin Online [132] Nov 2d 3d 1878 Tropaeolum majus Not young internodes on nor Plants were observed for the sake of expected apheliotropism, but they were Heliotropic. A plant 10 1/2 inches high had glass filament with 2 little triangles of paper attached to 4th internode from summit, (upper internode short short) the point of attachment of filament 8 1/2 inches beneath horizontal glass so tracing fairly much enlarged— Plant surrounded on all sides on summit with
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bowed growing old the next morning Dec. 30th 6° 50' a.m. The straight course pursued from 8° 53' to 12° I am inclined to attribute to t. being having in Hot-House become bowed towards light, when position of plant reversed, it travelled in straight line to light, afterwards circumnutated — From After 12° to I can see no signs of Heliotropism or Apheliotropism. Dec. 30 Two young tendrils were observed the Pot having been reversed compared to yesterd; they moved from the light till, in one case
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April 2d all shoots straight April 5th placed in light again — on 9th 2 of them beginning to be rectangular curve, on 11th one rectangular — On 18th put again in dark, plant with 6 shoots rectangular — on 20th greatly raised some almost vertical. 21' two uppermost almost completely vertical — 22 two others completely vertical — (I conclude that shoot grow out rectangularly by epinasty are kept down by by apheliotropism — They are, lower certainly apogeotropic [33v
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The Complete Work of Charles Darwin Online [65] N.B. Lathyrus aphaca — I can detect no apheliotropism in tendrils, yet they circumnutate very little — top of stem is highly heliotropic — Nor can I in Greenhouse see any signs in L. maritimus.— Sept 2d. 1879
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. (I suspect season of year) (N.B I suppose from effect in Hothouse that light fr N.E window not sufficient to induce apheliotropism— but it is very odd that both in study Hot-House the shoots were heliotropic) Give case of circumnutation under circumnutation of stems or under Stolons I have mentioned Trailing Branches Also worth giving under Heliotropism Apheliotropism— — when I mention about Tropaeolum at first heliotropic then said to be apheliotropi
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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the other seedling, traced under the same circumstances, were closely similar. Apheliotropism. We succeeded in observing only two cases of apheliotropism, for these are somewhat rare; and the movements are generally so slow that they would have been very troublesome to trace. Fig. 178. Bignonia capreolata: apheliotropic movement of a tendril, traced on a horizontal glass from 6.45 A.M. July 19th to 10 A.M. 20th. Movements as originally traced, little magnified, here reduced to two-thirds of the
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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CHAPTER VIII. MODIFIED CIRCUMNUTATION: MOVEMENTS EXCITED BY LIGHT. Distinction between heliotropism and the effects of light on the periodicity of the movements of leaves Heliotropic movements of Beta, Solanum, Zea, and Avena Heliotropic movements towards an obscure light in Apios, Brassica, Phalaris, Tropæolum, and Cassia Apheliotropic movements of tendrils of Bignonia Of flower-peduncles of Cyclamen Burying of the pods Heliotropism and apheliotropism modified forms of circumnutation Steps by
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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movement of circumnutation which gives rise to, or is converted into, heliotropism and apheliotropism. On this view we need not assume against all analogy that a lateral light entirely stops circumnutation; it merely excites the plant to modify its movement for a time in a beneficial manner. The existence of every possible gradation, between a straight course towards a lateral light and a course consisting of a series of loops or ellipses, becomes perfectly intelligible. Finally, the
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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) fixed transversely a little above the apex. It plainly circumnutated (Fig. 193) whilst increasing in length and growing downwards. It was then raised up, so as to be extended almost horizontally, and the terminal part curved itself downwards, following a nearly straight course during 12 h., but with one attempt to circumnutate, as shown in Fig. 194. After 24 h. it had become nearly vertical. Whether the exciting cause of the downward movement is geotropism or apheliotropism was not ascertained
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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Heliotropic movements of Beta, Solanum, Zea, and Avena Heliotropic movements towards an obscure light in Apios, Brassica, Phalaris, Tropæolum, and Cassia Apheliotropic movements of tendrils of Bignonia Of flower-peduncles of Cyclamen Burying of the pods Heliotropism and apheliotropism modified forms of circumnutation Steps by which one movement is converted into the other Transversal-heliotropismus or diaheliotropism influenced by epinasty, the weight of the part and apogeotropism Apogeotropism overcome
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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Lateral illumination of the tip determines the direction of the curvature of the base Cotyledons of Avena, curvature of basal part due to the illumination of upper part Similar results with the hypocotyls of Brassica and Beta Radicles of Sinapis apheliotropic, due to the sensitiveness of their tips Concluding remarks and summary of chapter Means by which circumnutation has been converted into heliotropism or apheliotropism Page 449-492 CHAPTER X. MODIFIED CIRCUMNUTATION: MOVEMENTS EXCITED BY
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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heliotropism, implies that a plant, when unequally illuminated on the two sides, bends from the light, instead of, as in the last sub-class of cases, towards it; but apheliotropism is comparatively rare, at least in a well-marked degree. There is a third and large sub-class of cases, namely, those of transversal-Heliotropismus of Frank, which we will here call diaheliotropism. Parts of plants, under this influence, place themselves more or less transversely to the direction whence the light
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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pot which had always stood upright, was left in the dark cupboard for six days; it bore 3 peduncles, and only one became within this time at all bowed downwards, and that doubtfully. The weight, therefore, of the pods is not the cause of the bending down. This pot was then brought back into the light, and after three days the peduncles were considerably bowed downwards. We are thus led to infer that the downward curvature is due to apheliotropism; though more trials ought to have been made. Fig
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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Sinapis apheliotropic, due to the sensitiveness of their tips Concluding remarks and summary of chapter Means by which circumnutation has been converted into heliotropism or apheliotropism. NO one can look at the plants growing on a bank or on the borders of a thick wood, and doubt that the young stems and leaves place themselves so that the leaves may be well illuminated. They are thus enabled to decompose carbonic acid. But the sheath-like cotyledons of some Gramineæ, for instance, those of
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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cut off for a length in three cases of .2 inch and in the four other cases of .14, .12, .1, and .07 inch. But these cotyledons, after being extended horizontally, bowed themselves upwards as effectually as the unmutilated specimens in the same pots, showing that sensitiveness to gravitation is not confined to their tips. GEOTROPISM. This movement is directly the reverse of apogeotropism. Many organs bend downwards through epinasty or apheliotropism or from their own weight; but we have met
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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radicles emit tertiary ones, but these, in the case of the bean, are not affected by gravitation; consequently they protrude in all directions. Thus the general * Dr. Karl Richter, who has especially attended to this subject ('K. Akad. der Wissenschaften in Wien,' 1879, p. 149), states that apheliotropism does not aid radicles in penetrating the ground. [page] 55
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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apheliotropism bending from the light. There is another reason for this change, for writers, as we have observed, occasionally drop the adjectives positive and negative, and thus introduce confusion into their discussions. Diaheliotropism may express a position more or less transverse to the light and induced by it. In like manner positive geotropism, or bending towards the centre of the earth, will be called by us geotropism; apogeotropism will mean bending in opposition to gravity or from the centre
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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was highly inclined, and the pot stood in front of a north-east window. During the five first days the frond moved downwards or became less inclined; and the long line which was traced was strongly zigzag, with loops occasionally formed or nearly formed; and this indicated circumnutation. Whether the sinking was due to epinastic growth, or apheliotropism, we do not know. As the sinking was slight on the fifth day, a new tracing was begun on the sixth day (Oct. 25th), and was continued for 47 h
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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upright, and when brought back into the light again became rectangularly curved, we believe that the bending is in part due to apheliotropism, apparently somewhat opposed by apogeotropism. On the other hand, from observing the effects of tying a shoot downwards, so that the rectangle faced upwards, we are led to believe that the curvature is partly due to epinasty. As the rectangularly bent portion of an upright stem grows older, the lower part straightens itself; and this is effected through
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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old enough, or the light was not sufficiently bright, to induce apheliotropism, for both plants bent slowly towards, instead of from the window during four days. The course, during two days of the first-mentioned internode, is given in Fig. 176; and we see that it either circumnutated on a small scale, or travelled in a zigzag line towards the light. We have thought this case of feeble heliotropism in one of the older internodes of a plant, [page] 43
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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half the cases their power of moving from the light. It is probable that if the tips had been cauterised for the length of a whole millimeter, all signs of apheliotropism would have disappeared. It may be suggested that although the application of caustic does not stop growth, yet enough may be absorbed to destroy the power of movement in the upper part; but this suggestion must be rejected, for we have seen and shall again see, that cauterising one side of the tip of various kinds of radicles
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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peduncles bend downwards, and this is due to epinasty; for on two occasions when pots were laid horizontally, the sub-peduncles assumed the same position relatively to the main peduncle, as would have been the case if they had remained upright; that is, each of them formed with it an angle of about 40 . If they had been acted on by geotropism or apheliotropism (for the plant was illuminated from above), they would have directed themselves to the centre of the earth. A main peduncle was secured
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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and from dependent branches, it cannot be due to apheliotropism or to epinasty, but must be attributed to geotropism. Nineteen * 'Hist. Phys. des Plantes d'Europe,' tom. ii. 1841, p. 106. [page] 51
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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ANODA. BRASSICA. Anoda Wrightii, sleep of cotyledons, 302, 312 , of leaves, 324 , downward movement of cotyledons, 444 Apheliotropism, or negative heliotropism, 5, 419, 432 Apios graveolens, heliotropic movements of hypocotyl, 422-424 tuberosa, vertical sinking of leaflets at night, 368 Apium graveolens, sleep of cotyledons, 305 , petroselinum, sleep of cotyledons, 304 Apogeotropic movements effected by joints or pulvini, 502 Apogeotropism, 5, 494; retarded by heliotropism, 501; concluding
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F1325
Book:
Darwin, C. R. 1880. The power of movement in plants. London: John Murray.
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, similarly secured to sticks, from the light, and although some of them rotted, many of their sub-peduncles turned very slowly from their reversed or from their horizontal positions, so as to stand in the normal manner parallel to the upper part of the main peduncle. These facts show that the movement is independent of geotropism or apheliotropism; it must therebe attributed to epinasty, which however is checked, at least as long as the flowers are young, by heliotropism. Most of the above flowers
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Ch X.) Burying of seed-capsules On the burying of seed-capsules.— One observation on this subject was made at a time when we cared were interested solely with the question how far various kinds kinds of movement consisted of modified circumnutation. We therefore neglected to observe determine, though this might easily have been done, without much difficulty, whether the downward movement by which certain capsules bury themselves was excited by geotropism or by apheliotropism. That it was not
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CUL-DAR226.1.130[.1]
Printed:
1863.06.13
Review of Movement in plants `Pharmaceutical Journal and Transactions': 486
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stems and tendrils, but those of positive and negative heliotropism, and of positive and negative geotropism—or, to adopt the phraseology in the present volume, of heliotropism, apheliotropism, geotropism and apogeotropism—all of which are but modifications of circumnutation. Probably the series of observations which will present the greatest novelty to most readers is that which demonstrates that these movements of nutation are as universal in the roots and other underground parts of plants as
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A268
Book:
Holder, Charles Frederick. 1892. Charles Darwin: his life and work. New York: G. P. Putnam's sons.
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the light; apheliotropism, or movement from the light; diaheliotropism or movement at right angles to the source of light; and paraheliotropism, embracing such movements as screen the plant from excess of light. To the second class belong: geotropism, ox movement towards the earth or into the soil; apogeotropism, or movement contrary to the force of gravity; and diageotropism, or movement at right angles to the force of gravity. The third class embraces the so-called nyctotropic movements of
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A297
Book:
Darwin, Francis & E. Hamilton Acton. 1909. Practical physiology of plants. Cambridge: University Press.
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. warmer than that in the vessel. The growth of the root will be at once accelerated, becoming nearly three times as quick. It will again fall as the water sinks to the temperature of the room. (184) Temperature: microscopic method. Repeat exp. 183, substituting water at 3° or 4° 0. for the water in which the observations were begun. Note the sudden and serious check to growth. 1 According to Kohl (Die ilechanik der Reizkriimmungen) bean seedlings grown on the klinostat show distinct apheliotropism
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A297
Book:
Darwin, Francis & E. Hamilton Acton. 1909. Practical physiology of plants. Cambridge: University Press.
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the leaves, may occur. The angles made by the leaves with the horizon should be noted, and should again be measured after a few days. (217) The movements due to specific sensitiveness. It was at one time believed that the diaheliotropic position was simply the result of a balance struck between such opposing tendencies as apheliotropism, apogeotropism, epinasty, c. c, and that diaheliotropism as a specific form of sensitiveness was non-existent. This view has now given way to the belief that a
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A297
Book:
Darwin, Francis & E. Hamilton Acton. 1909. Practical physiology of plants. Cambridge: University Press.
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water-culture, 162 Alkali, use of, in extraction, 245 Alkalinity, see Acids Amaranthus, anthocyan in, 52 Amides, qualitative tests for, 252 ; estimation of, 257, 258 ; experiments on, 259, 260 Ammonia, 253, 200, 315 ; estimation of, 259 Analysis, preparatory treatment of material for, 240 Anisotropism, 121, 122 Antherozoids attracted by malic acid, 214 Anthocyan, 52 Apheliotropism, 182 Apogeotropism, 164 Apparatus for culture of plants without C02, 28, 30 ; for recording geotropic and sleep
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A297
Book:
Darwin, Francis & E. Hamilton Acton. 1909. Practical physiology of plants. Cambridge: University Press.
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nitrogen applied to proteids, etc. 255 Kleinia, bloom of, 111 Klinostat, theory of action of, 186; use of in studying diaheliotro-pism, 186; used for exclusion of heliotropism and geotropism, 196; vertical, for growth experiments, 161 Knight's centrifugal experiment, 168 Enop on water culture, 59 n. Konig on oils and fats, 261 Kohl's method of recording absorption of transpiring plant, 83; on light affecting transpiration, 85, 85 n.; on apheliotropism of bean roots, 157 n. Krabbe on
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