Investigations of the Geotropic Curvature of the Avena Coleoptile

The geotropic reaction in Avena coleoptiles is studied as a function of the stimulation time. The direction of the stimulation with respect to the vascular bundles must be defined when studying geotropic responses. It is found that the threshold time to evoke geotropic response is less than half a minute, i.e., at least ten times lower than the presentation time usually reported in the literature. An extrapolation procedure can be used to give a so-called extrapolated presentation time t_b, which is intimately related to the logarithmic part of the geotropic response curve and has a physical meaning in the reciprocity rule. The problem of the duration of the true threshold time for stimulation with 1 g is discussed. An experiment indicates that it is not necessary for mass particles (“statoliths”) to settle on the lateral cell wall in order to start the geotropic reaction chain. The slope of the logarithmic part of the geotropic response curve is independent of the transverse force applied to the coleoptiles. Support is given to the view that the slope is determined by the number of sedimenting mass particles.     

Kinetics of stress relaxation properties of oat coleoptile cell wall after geotropic stimulation

This study describes the stress relaxation of the cell wall of oat (Avena sativa) coleoptiles after different periods of geotropic stimulation. The upper and lower tissues (with respect to gravity) of geotrophically stimulated coleoptiles exhibit different wall properties. The lower tissues are less resistant to deformation than the upper. The ratio of stress to strain is significatly less in the lower than in the upper tissue. Similarly, the relaxation time and the minimum relaxation time, derived from the Maxwell model which describes the physical characteristics of polymers, are also shorter in the lower tissue. However, the maximum relaxation time shows no difference between the upper and lower tissues of a geotropically stimulated coleoptile. The differences between the tissues begin at about 8 minutes after the commencement of stimulation, similar to the time for the initiation of dictyosome redistribution, and precede the onset of geotropism. The above responses of the cell wall of the lower tissue are similar to those induced by indoleacetic acid. The parameters of wall properties of the coleoptiles of both the control and the geostimulated fluctuate rhythmically with time. The periodic changes in wall properties of the coleoptile are compared to other cyclic physiological phenomena.     

Physiology of Morphactins: Effect on Gravi- and Photo-response

Plant roots and shoots respond to gravity and light source in a definite way. Thus, there are typical geotropic and phototropic responses for roots and shoots. When seedlings were grown in presence of morphactins, IT 3233 or IT 3456, on a vertical or a horizontal plane, the roots and shoots lost the capacity to respond to gravity or to unilateral light source. This was true for both monocots and dicots. This suggests that basic mechanism (s) of the two tropic responses are the same in the roots and shoots of the two plant groups. The site(s) of action of morphactins is unknown. The reaction (s) controlling geotropism and phototropism may be closely related as morphactins affected both geotropic and phototropic response of the same organ. Indoleacetic acid and gibberellic acid per se did not modify the effect of morphaclins on geotropism. Growth retardation effect of morphactins appears to be controlled by another mechanism.     

Geotropism and the lateral transport of auxin in the corn mutant amylomaize

Summary In coleoptiles of the amylomaize corn mutant (AM), the amyloplasts are much reduced in size in comparison with the wild type corn (WT), permitting a comparison of geotropic responsiveness as related to lateral displacement of amyloplasts and lateral transport of auxin. The amyloplasts of AM showed 30–40% lesser lateral redistribution in response to horizontal exposure in comparison with WT. With geotropic stimulation, the lateral transport of auxin in the direction of growth was 40–80% less, and the geotropic curvature by the coleoptiles was also significantly less in the mutant as compared with WT. These correlations support the hypothesis that the starch plastids serve as gravity sensors in the geotropic responses of coleoptiles.     

Wirkung von Vorbestrahlung mit Rot- oder Blaulicht auf die geotropische Empfindlichkeit von Maiskoleoptilen

The early geotropic downward bending of corn (Zea mays L.) coleoptiles was found to be influenced by red and blue light. The coleoptiles were illuminated from above and kept in the dark for defined intervals; afterwards they were positioned horizontally and their curvature was monitored for 40 min. After illumination with red light and a 120 min interval and early upward bending instead of an early downward bending was found. This effect was nullified by a far-red illumination administered immediately after exposure to red light. These results indicate that the phytochrome system influences the geotropic reaction. After illumination with blue light and a 30 min interval little downward bending was found. This result corresponds well with the findings of earlier authors who measured the late geotropic reaction, on the basis of the hypothesis that the strength of the early downward bending is a measure of the geotropic sensitivity. The dose-effect curve of the blue light influence on geotropic sensitivity, measured by early downward bending, is very similar to the dose-effect curve of the phototropic curvature of corn coleoptiles. This fact together with the earlier finding of similar adaptation times of about 30 min suggests the existence of some common transducers in the reaction chain of phototropism and geotropism.

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Abkürzungen HR Hellrot - DR Dunkelrot - D Dunkel - WT Wartezeit - DK Dunkelkontrolle     

The Effects of Red, Far-red, and Blue Light on the Geotropic Response of Coleoptiles of Zea mays

The effects of red, far-red, and blue light on the geotropicresponse of excised coleoptiles of Zea mays have been investigated.Seedlings were grown in darkness for 5 or 6 days, exposed tovarious light treatments, and then returned to darkness fordetermination of the geotropic response. The rate of response of the coleoptiles is decreased after theyhave been exposed to red light (620–700 mµ, 560ergs cm^–2sec^–1 for the 24 hrs, but not for the 4hrs, preceding stimulation by gravity. Furthermore, their rateof response is greatly reduced if they are exposed to red lightfor 10 min and then returned to darkness for 20 hrs before geotropicstimulation. At 25° C an interval of 6 to 8 hrs elapses between a 10-minexposure to red light and the first detectable decrease in thegeotropic response of the coleoptile. This interval can be lengthenedby exposing the seedlings to low temperatures (0° to 2°C) after the light treatment but cannot be greatly shortenedby increasing the duration of exposure to red light. Using a standard procedure of exposing 5-day-old etiolated seedlingsto light for various times, replacing them in darkness for 20hrs and then determining the response of the coleoptiles to4 hrs geotropic stimulation, it has been found that: (a) Exposureto red light for 15 sec significantly decreases the geotropiccurvature of the coleoptiles and that further reduction occurson increasing the length of the light treatment to 2 and 5 min.(b) Far-red light has no effect on the geotropic response ofthe coleoptiles but it can completely reverse the effect ofred light. After repeated alternate exposure to red and far-redlight the geotropic response of the coleoptile is determinedby the nature of the last exposure, (c) Complete reversal ofthe effect of red light by far-red radiation only occurs whenexposure to far-red follows immediately after exposure to red.The reversing effect of far-red radiation is reduced if a periodof darkness intervenes between the red and far-red light treatments,and is lost after a dark interval of approximately 2 hrs. The effect of red light on the rate of geotropic response ofthe coleoptiles is independent of their age and length at thetime of excision. Blue light acts in a similar way to red light, but the seedlingsare less sensitive to blue than to red light. Coleoptiles grown throughout in a mixture of continuous, weak,red, and far-red light have a lower rate of geotropic responsethan etiolated coleoptiles.     

Gravitropism of maize and rice coleoptiles: dependence on the stimulation angle

Gravitropism of maize and rice coleoptiles was investigated with respect to its dependence on the angle of displacement or the initial stimulation angle (ISA). Close examination of curvature kinetics and the response to a drop in stimulation angle (SA) indicated that the gravtropic response during an early but substantial part of the curvature development is directly related to the ISA, there being no effect of the reduction of SA resulting from the curvature response itself. On the basis of this finding, the relationship between the steady SA and the curvature rate was determined. In maize, the curvature rate increased linearly with the sines of SAs up to an SA of 90 degrees. Rice coleoptiles, however, showed a saturation curve in the same range of SAs. The saturation profile was nearly identical between coleoptiles grown in air and those submerged in water, although the latter elongated much faster. Rice coleoptiles appeared to be far more sensitive to gravity than maize coleoptiles. It is concluded that the sensitivity to gravity, assessed through dependence on ISA, is a property inherent to a given gravitropic organ. Long-term measurements of curvature indicated that the coleoptiles bend back past the vertical. This overshooting was marked in submerged rice coleoptiles.     

Orientation of wheat seedling organs in relation to gravity

Seedlings of wheat (Triticum aestivum L.) were grown in special holders that permitted the coleoptile and early roots to develop in moist air. The orientation of the organs of seedlings erect to gravity was compared with that of organs produced on a horizontal clinostat. Orientation was described by the angular position of each organ tip with reference to the axis of the embryo. Comparative tests were also made with barley, rye, and oat seedlings.

The coleoptile of all species developed curvatures in 3 dimensions when geotropic responses were eliminated. The primary root was not precise in its positive geotropism. Seedlings grew on clinostats with much greater variations in the lateral orientation of the central root and with a tendency for it to curve away from the endosperm to a greater degree than in erect seedlings.

The symmetry of root system in wheat was found to depend on a specific mechanism. Under the influence of gravity the earliest lateral roots were oriented in a plane at characteristic angles of about 57.5° with the ideal primary root. The corresponding angles for lateral roots growing on clinostats were greater by about 47.5° as a result of epinasty not previously reported in roots. This force also appeared to be active in the seminal roots of barley and rye but not of oats.

The curvatures in coleoptiles grown without the directional effects of gravity correspond to the results of growth imbalance in Coleus stems in the absence of lateral transport of their auxin by gravity. Root epinasty appears to be based on auxin imbalance. Curvatures in the primary root are also interpreted as results of asymmetrical distribution of growth hormone.

     

Early bending kinetics in response to geostimulation or unilateral IAA application in Lolium nodal segments

Abstract In Lolium multiflorum nodal segments, bending responses both to geostimulation and unilateral indole-3-acetic acid (IAA) application exhibited much variability in their lag times and speeds of early bending. Despite this variability, mean response curves to gravity and auxin stimulus were markedly similar with each having a phase of immediate, negative bending followed by phases of slow, positive bending and eventually more rapid, positive bending within 40 min of initial treatment. Comparison of lag times for response to geostimulation and unilateral IAA application, whether derived from the mean of individual replicates, or from mean curve data, showed that at least 4 min is available in this geotropic system for establishment of asymmetric auxin levels that could lead to differential growth. The hypothesis that variability in georesponse in Lolium nodal segments is linked to variable sensitivity of geosensitive tissue to auxin was tested using matching longitudinally-halved nodal segments and evidence was obtained in support of the hypothesis from lag time but not from early bending speed data. The implications of the findings for an involvement of endogenous IAA in shoot geotropism together with the necessity to understand better the complex behaviour of bending response in individual replicates are discussed.     

Gravitropic responses of the Avena coleoptile in space and on clinostats. I. Gravitropic response thresholds

We conducted a series of gravitropic experiments on Avena coleoptiles in the weightlessness environment of Spacelab. The purpose was to test the threshold stimulus, reciprocity rule and autotropic reactions to a range of g-force stimulations of different intensities and durations The tests avoided the potentially complicating effects of earth's gravity and the interference from clinostat ambiguities. Using slow-speed centrifuges, coleoptiles received transversal accelerations in the hypogravity range between 0.1 and 1.0 g over periods that ranged from 2 to 130 min. All responses that occurred in weightlessness were compared to clinostal experiments on earth using the same apparatus.
Characteristic gravitropistic response patterns of Avena were not substantially different from those observed in ground-based experiments. Gravitropic presentation times were extrapolated. The threshold at 1.0 g was less than 1 min (shortest stimulation time 2 min), in agreement with values obtained on the ground. The least stimulus tested, 0.1 g for 130 min, produced a significant response. Therefore the absolute threshold for a gravitropic response is less than 0.1 g.     

The lateral transport of IAA in intact coleoptiles of Avena sativa L. and Zea mays L. during geotropic stimulation

Summary Movement of IAA was studied in excised coleoptile apices and whole seedlings of Zea mays L. and Avena sativa L. during geotropic stimulation. A micropipette technique permitted the application of [5-³H]IAA at predetermined points on the coleoptiles with minimal tissue damage.When [5-³H]IAA was applied to the upper side of a horizontal excised Zea coleoptile, about 60% of the recoverable radioactivity had moved into the lower half after 2 h. In contrast, when application was made to the lower side of a horizontal excised coleoptile, only 4% of the radioactivity migrated to the upper half. There was, thus, a net downward movement of 56%. Similar patterns of distribution were found for radioactivity in both the tissue and the basal receiver blocks. In horizontal shoot tissues of intact Zea seedlings a net downward movement of about 30% of the recoverable radioactivity occurred after 1 h of geotropic stimulation. Comparable experiments with Avena indicated a net downward movement of 6–12% in excised apices of coleoptiles and in the intact shoot. In both Zea and Avena chromatographic analyses of tissue and receiver blocks indicated that the movement of radioactivity reflected that of IAA.In Zea coleoptiles, the lateral migration of radioactivity after 2 h was 3 to 4 times greater in the apical tissues than in the basal tissues. A significant net downward movement of radioactivity was detected after 10 min of geotropic stimulation in the extreme apex of Zea coleoptiles but not in the more basal regions.These experiments show that downward lateral transport of IAA occurs in intact shoots of Zea and Avena seedlings upon geotropic stimulation. Lateral transport of IAA had previously been demonstrated only in sub-apical segments of Zea coleoptiles.     

Enhancement of phototropic response to a range of light doses in Triticum aestivum coleoptiles in clinostat-simulated microgravity

The phototropic dose-response relationship has been determined for Triticum aestivum cv. Broom coleoptiles growing on a purpose-built clinostat apparatus providing gravity compensation by rotation about a horizontal axis at 2 rev·min^-1. These data are compared with data sets obtained with the clinostat axis vertical and stationary, as a 1·g control, and rotating vertically to examine clinostat effects other than gravity compensation. Triticum at 1·g follows the wellestablished pattern of other cereal coleoptiles with a first positive curvature at low doses, followed by an indifferent response region, and a second positive response at progressively increasing doses. However, these response regions lie at higher dose levels than reported for Avena. There is no significant difference between the responses observed with the clinostat axis vertical in the rotating and stationary modes, but gravity compensation by horizontal rotation increases the magnitude of first and second positive curvatures some threefold at 100 min after stimulation. The indifferent response is replaced by a significant curvature towards the light source, but remains apparent as a reduced curvature response at these dose levels.     

The Change of Tropism in Polyporus brumalis Stipes and the Effect of Directional Stimuli on Pileus Differentiation

The directional influences of light and gravity upon the developmentof pileate and epileate sporophores of Polyporus brumalis, producedin pure culture, have been investigated. Growth in the dimiticfruit-body has a considerable subapical component which in thestipe is responsible for tropistic curvature. The stipe is competentto react negatively to unilateral gravitational and positivelyto unilateral photic stimulation throughout development, butwhen both stimulioperate phototropism masks geotropism. If illuminatedfrom one side the growing epileate stipe is strongly positivelyphototropic but as the pileus reaches a characteristic diameterthe stipe becomes negatively geotropic. Experiments with changingdirection of illumination and with artificial pilei of blackpaper suggest that the change of tropism is explicable by theshading action of the expanding pileus on the sub-pilcal photoperceptiveand photoreactive region of the stipe. Developing sporophores continuously rotated with reference tofixed directions of gravitational and light stimulation andothers with stipes inverted as a result of illumination frombelow during development have in common that the morphologicallyupper surface of the pileus always develops towards and approximatelyat right angles to the direction from which the maximum lightintensity is received. Normal but inverted stipes and dissepimentsshow no tendency to geotropic reorientation.     

The Mechanism of Geoperception in Lentil Roots

The sediment of the amyloplasts in the statocytes of lentilroots was analysed in two different samples of seedlings. Theroots of the first sample (HP) were grown in a horizatal position,whereas thoes of the second sample (VP+20) were grown in a horizontalposition and then exposure to give a response). It is demonstratedthat the statolish can almost enter into contact with the plasmamembrane lining the longitudinal wall of the statocytes of theHP sample. However, these organelles in the VP+20 sample areable to provoke a geotropic stimulation though they are sedimentedat a distance of 0.1350–0.2600 µm from the plasmalemma.From these data it is concluded that the cytoplasmic structureswhich may play a role in the geotropic stimulation are: (1)the endoplasmic reticulum located along the longitudianl wallof the statocytes and (2) the microtubules or the plasmalemmaif the action of the statoliths is transmitted by the parietalcytoplasm. The large aggregations of endoplasmic reticulum whichare situated in the distal part of the central root cap cellswould not have any role in the perception of gravity by roots.These results are discussed in the light of recent work showingthe action of a growth inibitor in the geotropic reaction ofroots.     

Lazy rice in space: gravity regulates helical movement in plants

Circumnutation, the helical movement of organs, has been observed in diverse species of land plants. Whether circumnutation arises purely from internal growth oscillations or as a response to exogenous forces such as gravity is a subject of active debate. By observing rice seedlings grown under microgravity at the International Space Station (ISS) and analyzing the agravitropic lazy1 mutant, Kobayashi et al. (2019) propose gravity as the causal force that regulates circumnutation of rice coleoptiles.     

Intracellular distribution of mitochondria after geotropic stimulation of the oat coleoptile

The number of mitochondria is greater in the bottom than in the top of cells in geotropically stimulated oat (Avena sativa) coleoptiles. In the avascular tip and outer epidermis of subapical regions this difference occurs only in the lower tissues. These inequalities are found both in the KMnO₄⁻ and in the glutaraldehyde-fixed tissues; however, they are significant only in the former. Also, the number of mitochondria scored is consistently lower when the tissues were fixed in KMnO₄. These results suggest that mitochondria undergo a small degree of sedimentation after geostimulation, a redistribution reduced by the slower fixation with glutaraldehyde. Differences in mitochondrial number begin later than those in the amyloplast and the Golgi apparatus after geotropic stimulation. The cells in the avascular-tip region (a region having an important role in geotropism) have two to three times more mitochondria than the subapical cells.     

Cell elongation in the grass pulvinus in response to geotropic stimulation and auxin application

Summary Horizontally-placed segments of Avena sativa L. shoots show a negative geotropic response after a period of 30 min. This response is based on cell elongation on the lower side of the leaf-sheath base (pulvinus). Triticum aestivum L., Hordeum vulgare L. and Secale cereale L. also show geotropic responses that are similar to those in Avena shoots. The pulvinus is a highly specialized organ with radial symmetry and is made up of epidermal, vascular, parenchymatous and collenchymatous tissues. Statoliths, which are confined to parenchyma cells around the vascular bundles, sediment towards the gravitational field within 10–15 min of geotropic stimulation. Collenchymatous cells occur as prominent bundle caps, and in Avena, they occupy about 30% of the volume of the pulvinus. Geotropic stimulation causes a 3- to 5-fold increase in the length of the cells on the side nearest to the center of the gravitational field. Growth can also be initiated in vertically-held pulvini by the application of indole-3-acetic acid, 1-naphthaleneacetic acid or 2.4-dichlorophenoxyacetic acid. 2.3.5.-triiodobenzoic acid interferes with growth response produced by geotropic stimulation as well as with the response caused by auxin application. Gibberellic acid and kinetin have no visible effect on the growth of the pulvinus. Polarization microscopy shows a unique, non-uniform stretching of the elongating collenchymatous cells. Nonelongated collenchymatous cells appear uniformally anisotropic. After geotropic stimulation or auxin application, they appear alternately anisotropic and almost isotropic. Such a pattern of cell elongation is also observed in collenchyma cells of geotropically-stimulated shoots of Rumex acetosa L., a dicotyledon.Abbreviations 2.4-D 2.4-dichlorophenoxyacetic acid - GA₃ gibberellic acid - IAA indole-3-acetic acid - NAA l-naphthaleneacetic acid - TIBA 2.3.5-triiodobenzoic acid     

Geotropic curvature of decapitated Avena coleoptiles after application of tips of maize roots,tips of Avena coleoptiles,indoleacetic acid,or abscisic acid

Isolated Avena coleoptiles were decapitated at different distances from the tip and then placed horizontally, after which the geotropic curvature was measured. No geotropic curvature could be detected during the first 3 h. Later, upward curvature occurred which was found to depend inversely on the length of the decapitated tips. When the tips of maize roots or Avena coleoptiles were placed on the cut surface of decapitated Avena coleoptiles, the coleoptiles showed a significantly stronger upward curvature as compared to controls which had been provided with agar blocks on the cut surface. The same upward curvature was found with decapitated coleoptiles provided with agar blocks containing 10^-6 or 10^-7 M indoleacetic acid (IAA). After application of abscisic acid (ABA) at concentrations of 10^-6 and 10^-8 M to the decapitated coleoptiles, the curvature observed was not different from that of the controls; at higher concentrations of ABA the curvature was found to be lower than that of the controls. It is concluded that root tips secrete a substance which may replace the effect of IAA in coleoptiles. The results are discussed in view of the validity of the Cholodny-Went hypothesis for the geotropic reaction of roots.Abbreviations ABA abscisic acid - IAA 3-indoleacetic acid     

Entrainment of Geotropic Oscillations in Hypocotyls of Heliunthus annuus– An Experimental and Theoretical Investigation

A theory for the geotropic movements of the Heliunthus unnuus hypocotyl has been given earlier by Johnsson et ul. In the present paper this theory is tested by extensive experiments. The magnitude as well as the rate of geotropic curvature are found to be in agreement with the theoretical predictions. Further, the geotropic reaction time, defined as the time between the start of geotropic stimulation and the start of the geotropic reaction is determined as a function of the stimulation angle with respect to the plumb line. A slight variation found in the reaction time is included in the theory.