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At least 109 records · Page 6

Effects of inversion on plastid position and gravitropism in Ceratodon protonemata

When dark-grown tip cells of protonemata of the moss Ceratodon purpureus are turned to the horizontal, plastids first sediment towards gravity in a specific zone and then the tip curves upward. To determine whether gravitropism and plastid sedimentation occur in other orientations, protonemata were reoriented to angles other than 90 degrees. Qualitative and quantitative light microscopic observations show that plastid sedimentation along the cell axis occurs in both upright and inverted cells. However, only some plastids fall and sedimentation is incomplete; plastids remain distributed throughout the length of the cell, and those plastids that sediment do not fall all the way to the bottom of the cell. Tip cells are gravitropic regardless of stimulation angle, and as in higher plants, the maximal rate of initial curvature is in response to a 120 degrees reorientation. Infrared videomicroscopy, time-lapse studies of living, inverted protonemata indicate that amyloplast sedimentation precedes upward curvature. Together, these data further support (i) the hypothesis that amyloplast sedimentation functions in gravitropic sensing in these cells, and (ii) the idea that gravity affected the evolution of cell organization.

NASA Program Space Biology↗

Microsurgical removal of epidermal and cortical cells: evidence that the gravitropic signal moves through the outer cell layers in primary roots of maize

There is general agreement that during root gravitropism some sort of growth-modifying signal moves from the cap to the elongation zone and that this signal ultimately induces the curvature that leads to reorientation of the root. However, there is disagreement regarding both the nature of the signal and the pathway of its movement from the root cap to the elongation zone. We examined the pathway of movement by testing gravitropism in primary roots of maize (Zea mays L.) from which narrow (0.5 mm) rings of epidermal and cortical tissue were surgically removed from various positions within the elongation zone. When roots were girdled in the apical part of the elongation zone gravitropic curvature occurred apical to the girdle but not basal to the girdle. Filling the girdle with agar allowed curvature basal to the girdle to occur. Shallow girdles, in which only two or three cell layers (epidermis plus one or two cortical cell layers) were removed, prevented or greatly delayed gravitropic curvature basal to the girdle. The results indicate that the gravitropic signal moves basipetally through the outermost cell layers, perhaps through the epidermis itself.

NASA Discipline Number 29-20↗

The role of extracellular free-calcium gradients in gravitropic signalling in maize roots

Gravitropism in roots has been proposed to depend on a downward redistribution of calcium across the root cap. However, because of the many calcium-binding sites in the apoplast, redistribution might not result in a physiologically effective change in the apoplasmic calcium activity. To test whether there is such a change, we measured the effect of gravistimulation on the calcium activity of statocyte cell walls with calcium-specific microelectrodes. Such a measurement must be made on a tissue with gravity sensing cells at the surface. To obtain such a tissue, decapped maize roots (Zea mays L. cv. Golden Cross Bantam) were grown for 31 h to regenerate gravitropic sensitivity, but not root caps. The calcium activity in the apoplasm surrounding the gravity-sensing cells could then be measured. The initial pCa was 2.60 +/- 0.28 (approx 2.5 mM). The calcium activity on the upper side of the root tip remained constant for 10 min after gravistimulation, then decreased 1.7-fold. On the lower side, after a similar lag the calcium activity increased 1.6-fold. Control roots, which were decapped but measured before recovering gravisensitivity (19 h), showed no change in calcium activity. To test whether this gradient is necessary for gravitropic curvature, we eliminated the calcium activity gradient during gravitropism by applying a mobile calcium-binding site (dinitro-BAPTA; 1,2-bis(2-amino-5-nitro-phenoxy)ethane-N,N,N',N'-tetraacetic acid) to the root cap; this treatment eliminated gravicurvature. A calcium gradient may be formed by proton-induced calcium desorption if there is a proton gradient. Preventing the formation of apoplastic pH gradients, using 10 and 50 mM 2-(N-morpholino)ethanesulfonic acid (Mes) buffer or 10 mM fusicoccin to stimulate proton excretion maximally, did not inhibit curvature; therefore the calcium gradient is not a secondary effect of a proton gradient. We have found a distinct and rapid differential in the apoplasmic calcium activity between the upper and lower sides of gravistimulated maize root tips which is necessary for gravitropism.

NASA Discipline Number 40-50↗

Root growth regulation and gravitropism in maize roots does not require the epidermis

We have earlier published observations showing that endogenous alterations in growth rate during gravitropism in maize roots (Zea mays L.) are unaffected by the orientation of cuts which remove epidermal and cortical tissue in the growing zone (Bjorkman and Cleland, 1988, Planta 176, 513-518). We concluded that the epidermis and cortex are not essential for transporting a growth-regulating signal in gravitropism or straight growth, nor for regulating the rate of tissue expansion. This conclusion has been challenged by Yang et al. (1990, Planta 180, 530-536), who contend that a shallow girdle around the entire perimeter of the root blocks gravitropic curvature and that this inhibition is the result of a requirement for epidermal cells to transport the growth-regulating signal. In this paper we demonstrate that the entire epidermis can be removed without blocking gravitropic curvature and show that the position of narrow girdles does not affect the location of curvature. We therefore conclude that the epidermis is not required for transport of a growth-regulating substance from the root cap to the growing zone, nor does it regulate the growth rate of the elongating zone of roots.

NASA Program Space Biology↗

Characterization of thermotropism in primary roots of maize: dependence on temperature and temperature gradient, and interaction with gravitropism

Thermotropism in primary roots of Zea mays L. was studied with respect to gradient strength (degrees C cm-1), temperature of exposure within a gradient, pre-treatment temperature, and gravitropic stimulation. The magnitude of the response decreased with gradient strength. Maximum thermotropism was independent of gradient strength and pre-treatment temperature. The range of temperature for positive and negative thermotropism did not change with pre-treatment temperature. However, the exact range of temperatures for positive and negative thermotropism varied with gradient strengths. In general, temperatures of exposure lower than 25 degrees C resulted in positive tropic responses while temperatures of exposure of 39 degrees C or more resulted in negative tropic responses. Thermotropism was shown to modify and reverse the normal gravitropic curvature of a horizontal root when thermal gradients were applied opposite the 1 g vector. It is concluded that root thermotropism is a consequence of thermal sensing and that the curvature of the primary root results from the interaction of the thermal and gravitational sensing systems.

Non-NASA Center↗

Hydrotropism and its interaction with gravitropism in maize roots

We have partially characterized root hydrotropism and its interaction with gravitropism in maize (Zea mays L.). Roots of Golden Cross Bantam 70, which require light for orthogravitropism, showed positive hydrotropism; bending upward when placed horizontally below a hydrostimulant (moist cheesecloth) in 85% relative humidity (RH) and in total darkness. However, the light-exposed roots of Golden Cross Bantam 70 or roots of a normal maize cultivar, Burpee Snow Cross, showed positive gravitropism under the same conditions; bending downward when placed horizontally below the hydrostimulant in 85% RH. Light-exposed roots of Golden Cross Bantam 70 placed at 70 degrees below the horizontal plane responded positively hydrotropically, but gravitropism overcame the hydrotropism when the roots were placed at 45 degrees below the horizontal. Roots placed vertically with the tip down in 85% RH bent to the side toward the hydrostimulant in both cultivars, and light conditions did not affect the response. Such vertical roots did not respond when the humidity was maintained near saturation. These results suggest that hydrotropic and gravitropic responses interact with one another depending on the intensity of one or both factors. Removal of the approximately 1.5 millimeter root tip blocked both hydrotropic and gravitropic responses in the two cultivars. However, removal of visible root tip mucilage did not affect hydrotropism or gravitropism in either cultivar.

NASA Program Space Biology↗

Correlations between gravitropic curvature and auxin movement across gravistimulated roots of Zea mays

We compared the kinetics of auxin redistribution across the caps of primary roots of 2-day-old maize (Zea mays, cv Merit) seedlings with the time course of gravitropic curvature. [3H] indoleacetic acid was applied to one side of the cap in an agar donor and radioactivity moving across the cap was collected in an agar receiver applied to the opposite side. Upon gravistimulation the roots first curved upward slightly, then returned to the horizontal and began curving downward, reaching a final angle of about 67 degrees. Movement of label across the caps of gravistimulated roots was asymmetric with preferential downward movement (ratio downward/upward = ca. 1.6, radioactivity collected during the 90 min following beginning of gravistimulation). There was a close correlation between the development of asymmetric auxin movement across the root cap and the rate of curvature, with both values increasing to a maximum and then declining as the roots approached the final angle of curvature. In roots preadapted to gravity (alternate brief stimulation on opposite flanks over a period of 1 hour) the initial phase of upward curvature was eliminated and downward bending began earlier than for controls. The correlation between asymmetric auxin movement and the kinetics of curvature also held in comparisons between control and preadapted roots. Both downward auxin transport asymmetry and downward curvature occurred earlier in preadapted roots than in controls. These findings are consistent with suggestions that the root cap is not only the site of perception but also the location of the initial redistribution of effectors that ultimately leads to curvature.

NASA Discipline Number 29-20↗

Organization of cortical microtubules in graviresponding maize roots

Immunofluorescence labeling of cortical microtubules (MTs) was used to investigate the relationship between MT arrangement and changes in growth rate of the upper and lower sides of horizontally placed roots of maize (Zea mays L. cv. Merit). Cap cells and cells of the elongation zone of roots grown vertically in light or darkness showed MT arrangements that were transverse (perpendicular) to the growth direction. Microtubules of cells basal to the elongation zone typically showed oblique orientation. Two hours after horizontal reorientation, cap cells of gravicompetent, light-grown and curving roots contained MTs parallel to the gravity vector. The MT arrangement on the upper side of the elongation zone remained transverse but the MTs of the outer four to five layers of cortical cells along the lower side of the elongation zone showed reorientation parallel to the axis of the root. The MTs of the lower epidermis retained their transverse orientation. Dark-grown roots did not curve and did not show reorientation of MTs in cells of the root cap or elongation zone. The data indicate that MT depolymerization and reorientation is correlated with reduction in growth rate, and that MT reorientation is one of the steps of growth control of graviresponding roots.

NASA Discipline Plant Biology↗

The mechanosensory calcium-selective ion channel: key component of a plasmalemmal control centre?

Mechanosensory calcium-selective ion channels probably serve to detect not only mechanical stress but also electrical, thermal, and diverse chemical stimuli. Because all stimuli result in a common output, most notably a shift in second messenger calcium concentration, the channels are presumed to serve as signal integrators. Further, insofar as second messenger calcium in turn gives rise to mechanical, electrical, and diverse chemical changes, the channels are postulated to initiate regulatory feedbacks. It is proposed that the channels and the feedback loops play a wide range of roles in regulating normal plant function, as well as in mediating disturbance of normal function by environmental stressors and various pathogens. In developing evidence for the physiological performance of the channel, a model for a cluster of regulatory plasmalemmal proteins and cytoskeletal elements grouped around a set of wall-to-membrane and transmembrane linkers has proved useful. An illustration of how the model might operate is presented. It is founded on the demonstration that several xenobiotics interfere both with normal channel behaviour and with gravitropic reception. Accordingly, the first part of the illustration deals with how the channels and the control system within which they putatively operate might initiate gravitropism. Assuming that gravitropism is an asymmetric expression of growth, the activities of the channels and the plasmalemmal control system are extrapolated to account for regulation of both rate and allometry of cell expansion. Finally, it is discussed how light, hormones, redox agents and herbicides could in principle affect growth via the putative plasmalemmal control cluster or centre.

NASA Program Space Biology↗

Introduction: an overview of gravity sensing, perception, and signal transduction in animals and plants

The antiquity of biological sensitivity and response to gravity can be traced through the ubiquity of morphology, mechanisms, and cellular events in gravity sensing biological systems in the most diverse species of both plants and animals. Further, when we examine organisms at the cellular level to elucidate the molecular mechanism by which a gravitational signal is transduced into a biochemical response, the distinction between plants and animals becomes blurred.

NASA Center HQS↗

Stimulation of root elongation and curvature by calcium

Ca2+ has been proposed to mediate inhibition of root elongation. However, exogenous Ca2+ at 10 or 20 millimolar, applied directly to the root cap, significantly stimulated root elongation in pea (Pisum sativum L.) and corn (Zea mays L.) seedlings. Furthermore, Ca2+ at 1 to 20 millimolar, applied unilaterally to the caps of Alaska pea roots, caused root curvature away from the Ca2+ source, which was caused by an acceleration of elongation growth on the convex side (Ca2+ side) of the roots. Roots of an agravitropic pea mutant, ageotropum, responded to a greater extent. Roots of Merit and Silver Queen corn also responded to Ca2+ in similar ways but required a higher Ca2+ concentration than that of pea roots. Roots of all other cultivars tested (additional four cultivars of pea and one of corn) curved away from the unilateral Ca2+ source as well. The Ca(2+)-stimulated curvature was substantially enhanced by light. A Ca2+ ionophore, A23187, at 20 micromolar or abscisic acid at 0.1 to 100 micromolar partially substituted for the light effect and enhanced the Ca(2+)-stimulated curvature in the dark. Unilateral application of Ca2+ to the elongation zone of intact roots or to the cut end of detipped roots caused either no curvature or very slight curvature toward the Ca2+. Thus, Ca2+ action on root elongation differs depending on its site of application. The stimulatory action of Ca2+ may involve an elevation of cytoplasmic Ca2+ in root cap cells and may partipate in root tropisms.

NASA Program Space Biology↗

Induction of curvature in maize roots by calcium or by thigmostimulation: role of the postmitotic isodiametric growth zone

We examined the response of primary roots of maize (Zea mays L. cv Merit) to unilateral application of calcium with particular attention to the site of application, the dependence on growth rate, and possible contributions of thigmotropic stimulation during application. Unilateral application of agar to the root cap induced negative curvature whether or not the agar contained calcium. This apparent thigmotropic response was enhanced by including calcium in the agar. Curvature away from objects applied unilaterally to the extreme root tip occurred both in intact and detipped roots. When agar containing calcium chloride was applied to one side of the postmitotic isodiametric growth zone ( a region between the apical meristem and the elongation zone), the root curved toward the side of application. This response could not be induced by plain agar. We conclude that curvature away from calcium applied to the root tip results from a thigmotropic response to stimulation during application. In contrast, curvature toward the calcium applied to the postmitotic isodiametric growth zone results from direct calcium-induced inhibition of growth.

NASA Discipline Number 40-50↗

The habits of roots: what's up down under?

Defining interactions of roots with the surrounding soil environment has been the focus of many recent investigations. As a result of these efforts, we are gaining an appreciation of the varied and often surprising strategies whereby roots adjust to and condition their soil environment for optimal growth and development. This article summarizes current knowledge of the often complex interactions between roots and biotic and abiotic factors within the soil. These interactions are interpreted in terms of modifications in the development or the physiology of the root.

Review↗

Cortical and cap sedimentation in gravitropic Equisetum roots

Although the rootcap is required for gravitropic sensing, various classical and contemporary data raise the question of whether additional sensing occurs away from the cap in roots. Roots of Equisetum hyemale L. (horsetail) were examined by light and electron microscopy to determine which cell components were distributed with respect to gravity both in and away from the rootcap. Adventitious roots from stem cuttings were gravitropic in a vertical orientation or if reoriented to the horizontal. Obvious amyloplast sedimentation was found in vertical and in reoriented roots 1) in cells in the center of the rootcap and 2) in young, elongating cortical cells located in two to three layers outside the endodermis. These cortical amyloplasts were smaller than cap amyloplasts and, unlike central cap amyloplasts, were occasionally found in the top of the cell. The nucleus was also sedimented on top of the amyloplasts in both cell types, both in vertical and in reoriented roots. Sedimentation of both organelles ceased as cortical cells elongated further or as cap cells became peripheral in location. In both cell types with sedimentation, endoplasmic reticulum was located in the cell periphery, but showed no obvious enrichment near the lower part of the cell in vertical roots. This is the first modern report of sedimentation away from the cap in roots, and it provides structural evidence that gravitropic sensing may not be confined to the cap in all roots.

NASA Discipline Plant Biology↗

Abscisic acid is not necessary for gravitropism in primary roots of Zea mays

Primary roots of Zea mays L. cv. Tx 5855 treated with fluridone are strongly graviresponsive, but have undetectable levels of abscisic acid (ABA). Primary roots of the carotenoid-deficient w-3, vp-5, and vp-7 mutants of Z. mays are also graviresponsive despite having undetectable amounts of ABA. Graviresponsive roots of untreated and wild-type seedlings contain 286 to 317 ng ABA g-1 f. wt, respectively. These results indicate that ABA is not necessary for root gravicurvature.

NASA Discipline Plant Biology↗

Defective secretion of mucilage is the cellular basis for agravitropism in primary roots of Zea mays cv. Ageotropic

Root caps of primary, secondary, and seminal roots of Z. mays cv. Kys secrete large amounts of mucilage and are in close contact with the root all along the root apex. These roots are strongly graviresponsive. Secondary and seminal roots of Z. mays cv. Ageotropic are also strongly graviresponsive. Similarly, their caps secrete mucilage and closely appress the root all along the root apex. However, primary roots of Z. mays cv. Ageotropic are non-responsive to gravity. Their caps secrete negligible amounts of mucilage and contact the root only at the extreme apex of the root along the calyptrogen. These roots become graviresponsive when their tips are coated with mucilage or mucilage-like materials. Peripheral cells of root caps of roots of Z. mays cv. Kys contain many dictyosomes associated with vesicles that migrate to and fuse with the plasmalemma. Root-cap cells of secondary and seminal (i.e. graviresponsive) roots of Z. mays cv. Ageotropic are similar to those of primary roots of Z. mays cv. Kys. However, root-cap cells of primary (i.e. non-graviresponsive) roots of Z. mays cv. Ageotropic have distended dictyosomal cisternae filled with an electron-dense, granular material. Large vesicles full of this material populate the cells and apparently do not fuse with the plasmalemma. Taken together, these results suggest that non-graviresponsiveness of primary roots of Z. mays cv. Ageotropic results from the lack of apoplastic continuity between the root and the periphery of the root cap. This is a result of negligible secretion of mucilage by cells along the edge of the root cap which, in turn, appears to be due to the malfunctioning of dictyosomes in these cells.

NASA Discipline Number 29-20↗

Influence of electrical fields and asymmetric application of mucilage on curvature of primary roots of Zea mays

Primary roots of Zea mays cv. Yellow Dent growing in an electric field curve towards the anode. Roots treated with EDTA and growing in electric field do not curve. When root cap mucilage is applied asymmetrically to tips of vertically-oriented roots, the roots curve toward the mucilage. Roots treated with EDTA curve toward the side receiving mucilage and toward blocks containing 10 mM CaCl2, but not toward "empty" agar blocks or the cut surfaces of severed root tips. These results suggest that 1) free calcium (Ca) is necessary for root electrotropism, 2) mucilage contains effector(s) that induce gravitropiclike curvature, and 3) mucilage can replace gravitropic effectors chelated by EDTA. These results are consistent with the hypothesis that the downward movement of gravitropic effectors to the lower sides of tips of horizontally-oriented roots occurs at least partially in the apoplast.

NASA Discipline Number 29-20↗