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At least 145 records · Page 8

Computer-assisted image analysis of plant growth, thigmomorphogenesis and gravitropism

A nonintrusive auxonometric system, based on the DARWIN image processor (Telewski et al. 1983 Plant Physiol 72: 177-181), is described and demonstrated in the analysis of gravitropism and thigmomorphogenesis in corn seedlings (Zea mays). Using this system, growth and bending of regularly shaped plants or organs can be quickly and accurately measured without, in any way, interfering with the plant. Furthermore, the growth and bending curves are automatically plotted. Thigmomorphogenesis in the aerial part of corn seedlings involves growth promotion at a low force load and growth retardation at higher force loads. The time courses of the two kinds of response are somewhat different, with retardation occurring immediately after mechanical perturbation and growth promotion taking somewhat longer to begin. Gravitropic experiments show that when dark-grown corn seedlings are placed on their side in the light, the resulting curvature is due to two consecutive morphological mechanisms. In the first instance, lasting for about 15 minutes, the elongation of the bottom edge of the plant accelerates, while the elongation of the top edge remains constant. After that, for the next 1.75 hours, the elongation of the top edge decelerates and stops while that of the bottom edge remains constant at the increased rate for most of the period. The measurements taken from both experiments at relatively high resolution (0.08-0.1 millimeter) show that the growth curves are not smooth but show many small irregularities which may or may not involve micronutations.

Zea mays/growth & development/physiology↗

Root gravitropism: a complex response to a simple stimulus?

Roots avoid depleting their immediate environment of essential nutrients by continuous growth. Root growth is directed by environmental cues, including gravity. Gravity sensing occurs mainly in the columella cells of the root cap. Upon reorientation within the gravity field, the root-cap amyloplasts sediment, generating a physiological signal that promotes the development of a curvature at the root elongation zones. Recent molecular genetic studies in Arabidopsis have allowed the identification of genes that play important roles in root gravitropism. Among them, the ARG1 gene encodes a DnaJ-like protein involved in gravity signal transduction, whereas the AUX1 and AGR1 genes encode proteins involved in polar auxin transport. These studies have important implications for understanding the intra- and inter-cellular signaling processes that underlie root gravitropism.

Review, Tutorial↗

High-resolution mapping and genetic characterization of the Lazy-2 gravitropic mutant of tomato

Mutation of the Lazy-2 (Lz-2) gene in tomato (Lycopersicon esculentum mill.) produces a phytochrome-dependent reversal of shoot gravitropism, providing a unique genetic resource for investigating how signals from light modulate gravitropism. We mapped the Lz-2 gene using RFLPs and a PCR-based technique to assess the feasibility of positional cloning. Analysis of a 1338 plant backcross population between L. esculentum and L. pennellii placed Lz-2 within a 1.2 cM interval on chromosome 5, 0.4 cM from TG504-CT201A interval. The inabililty to resolve these markers indicates that Lz-2 resides in a centromeric region in which recombination is highly suppressed. Lazy-2 is tightly linked to but does not encode the gene for ACC4, an enzyme involved in ethylene biosynthesis. We also observed that Lz-2 is partially dominant under certain conditions and stages of development.

Non-NASA Center↗

Enhanced Labeling Techniques to Study the Cytoskeleton During Root Growth and Gravitropism

Gravity effects the growth and development of all living organisms. One of the most obvious manifestations of gravity's effects on biological systems lies in the ability of plants to direct their growth along a path that is dictated by the gravity vector (called gravitropism). When positioned horizontally, in florescence stems and hypocotyls in dicots, and pulvini in monocots, respond by bending upward whereas roots typically bend downward. Gravitropism allows plants to readjust their growth to maximize light absorption for photosynthesis and to more efficiently acquire water and nutrients form the soil. Despite its significance for plant survival, there are still major gaps in understanding the cellular and molecular processes by which plants respond to gravity. The major aim of this proposal was to develop improved fluorescence labeling techniques to aid in understanding how the cytoskeleton modulated plant responses to gravity.

Blancaflor, Elison B.↗

Control of gravitropic orientation. I. Non-vertical orientation by primary roots of maize results from decay of competence for orthogravitropic induction

Plant organs may respond to gravity by vertical (orthogravitropic), oblique (plagiogravitropic) or horizontal (diagravitropic) growth. Primary roots of maize (Zea mays L.) provide a good system for studying such behaviours because they are reportedly capable of displaying all three responses. In current work using maize seedlings of the Silver Queen cultivar, stabilisation of growth at an oblique orientation was commonplace. Hypothetically, plagiogravitropism may be accomplished either by a process we call graded orthogravitropism or by hunting about a sensed non-vertical setpoint. In graded orthotropism primary bending is unidirectional and depends on facilitative stimuli that determine its extent. The hallmark of the setpoint mechanism is restorative curvature of either sign following a displacement; both diagravitropism and orthogravitropism are based on setpoints. Roots settled in a plagiogravitropic orientation were tested with various illumination and displacement protocols designed to distinguish between these two hypotheses. The tests refuted the setpoint hypothesis and supported that of graded orthotropism. No evidence of diagravitropism could be found, thus, earlier claims were likely based on inadequately controlled observations of graded orthotropism. We propose that orthotropism is graded by the sequential action of dual gravity receptors: induction of a vectorial gravitropic response requires gravitational induction of a separate facilitative response, whose decay in the absence of fresh stimuli can brake gravitropism at plagiotropic angles.

NASA Discipline Plant Biology↗

Wall extensibility and gravitropic curvature of sunflower hypocotyls: correlation between timing of curvature and changes in extensibility

Gravitropic curvature results from unequal growth rates on the upper and lower sides of horizontal stems. These unequal growth rates could be due to differences in wall extensibility between the two sides. To test this, the time course of curvature of horizontal sunflower (Helianthus annuus L.) hypocotyls was determined and compared with the time courses of changes in Instron-measured wall extensibility (PEx) of the upper and lower epidermal layers. As gravicurvature developed, so did the difference in PEx between the upper and lower epidermis. The enhanced growth rate on the lower side during the period of maximum increase in curvature was matched by PEx values greater than those of the vertical control, while the inhibited growth rate on the upper side was accompanied by PEx values below that of the control. The close correlation between changes in growth rates and alterations in PEx demonstrates that changes in wall extensibility play a major role in controlling gravicurvature.

Non-NASA Center↗

Basipetal auxin transport is required for gravitropism in roots of Arabidopsis

Auxin transport has been reported to occur in two distinct polarities, acropetally and basipetally, in two different root tissues. The goals of this study were to determine whether both polarities of indole-3-acetic acid (IAA) transport occur in roots of Arabidopsis and to determine which polarity controls the gravity response. Global application of the auxin transport inhibitor naphthylphthalamic acid (NPA) to roots blocked the gravity response, root waving, and root elongation. Immediately after the application of NPA, the root gravity response was completely blocked, as measured by an automated video digitizer. Basipetal [(3)H]IAA transport in Arabidopsis roots was inhibited by NPA, whereas the movement of [(14)C]benzoic acid was not affected. Inhibition of basipetal IAA transport by local application of NPA blocked the gravity response. Inhibition of acropetal IAA transport by application of NPA at the root-shoot junction only partially reduced the gravity response at high NPA concentrations. Excised root tips, which do not receive auxin from the shoot, exhibited a normal response to gravity. The Arabidopsis mutant eir1, which has agravitropic roots, exhibited reduced basipetal IAA transport but wild-type levels of acropetal IAA transport. These results support the hypothesis that basipetally transported IAA controls root gravitropism in Arabidopsis.

Non-NASA Center↗

Gravitropism of hypocotyls of wild-type and starch-deficient Arabidopsis seedlings in spaceflight studies

The major purpose of this spaceflight project was to investigate the starch-statolith hypothesis for gravity perception, and a secondary goal was to study plant growth and development under spaceflight conditions. This research was based on our ground studies of gravity perception in the wild type and three starch-deficient (one starchless and two reduced starch) mutants of Arabidopsis thaliana (L.) Heynh. Dark-grown seedlings that developed in microgravity were given one of several (30 min, 60 min, or 90 min) 1-g stimuli by an on-board centrifuge, and additional controls for seedling development also were performed. These latter control experiments included a morphological study of plants that developed in space in microgravity (F microg), in space on a centrifuge (F 1g), on the ground (G 1g), and on a rotating clinostat on the ground. Since elevated levels of ethylene were reported in the spacecraft atmosphere, additional controls for morphology and gravitropism with added ethylene also were performed. While exogenous ethylene reduced the absolute magnitude of the response in all four strains of Arabidopsis, this gas did not appear to change the relative graviresponsiveness among the strains. The relative response of hypocotyls of microgravity-grown seedlings to the stimuli provided by the in-flight centrifuge was: wild type > starch-deficient mutants. Although the protoplast pressure model for gravity perception cannot be excluded, these results are consistent with a statolith-based model for perception in plants.

Non-NASA Center↗

Root gravitropism

When a plant root is reoriented within the gravity field, it responds by initiating a curvature which eventually results in vertical growth. Gravity sensing occurs primarily in the root tip. It may involve amyloplast sedimentation in the columella cells of the root cap, or the detection of forces exerted by the mass of the protoplast on opposite sides of its cell wall. Gravisensing activates a signal transduction cascade which results in the asymmetric redistribution of auxin and apoplastic Ca2+ across the root tip, with accumulation at the bottom side. The resulting lateral asymmetry in Ca2+ and auxin concentration is probably transmitted to the elongation zone where differential cellular elongation occurs until the tip resumes vertical growth. The Cholodny-Went theory proposes that gravity-induced auxin redistribution across a gravistimulated plant organ is responsible for the gravitropic response. However, recent data indicate that the gravity-induced reorientation is more complex, involving both auxin gradient-dependent and auxin gradient-independent events.

NASA Discipline Plant Biology↗

Gravitropic moss cells default to spiral growth on the clinostat and in microgravity during spaceflight

In addition to shoots and roots, the gravity (g)-vector orients the growth of specialized cells such as the apical cell of dark-grown moss protonemata. Each apical cell of the moss Ceratodon purpureus senses the g-vector and adjusts polar growth accordingly producing entire cultures of upright protonemata (negative gravitropism). The effect of withdrawing a constant gravity stimulus on moss growth was studied on two NASA Space Shuttle (STS) missions as well as during clinostat rotation on earth. Cultures grown in microgravity (spaceflight) on the STS-87 mission exhibited two successive phases of non-random growth and patterning, a radial outgrowth followed by the formation of net clockwise spiral growth. Also, cultures pre-aligned by unilateral light developed clockwise hooks during the subsequent dark period. The second spaceflight experiment flew on STS-107 which disintegrated during its descent on 1 February 2003. However, most of the moss experimental hardware was recovered on the ground, and most cultures, which had been chemically fixed during spaceflight, were retrieved. Almost all intact STS-107 cultures displayed strong spiral growth. Non-random culture growth including clockwise spiral growth was also observed after clinostat rotation. Together these data demonstrate the existence of default non-random growth patterns that develop at a population level in microgravity, a response that must normally be overridden and masked by a constant g-vector on earth.

Non-NASA Center↗

The mechanics of gravitropic bending in leafy dicot stems

The mechanism of the gravitropic bending in stems of the cocklebur and castor bean are investigated. The results of these experiments demonstrate the quick stopping of growth and the increased tensions on the upper layer of a horizontal stem. It is suggested that bending apparently occurs as the resistance of the upper surface layers is extended to the inner cells below. A model of stem bending is developed which can explain the asymmetry of the stem-cell response.

Salisbury, F. B.↗

Gravitropic basis of leaf blade nastic curvatures

The curvatures produced in leaf blades by auxin treatment have been described as nastic curvatures because the initial differential growth is always enhanced on the lower side, regardless of the side of application. It is now known, however, that blades can show differential growth of either the upper or the lower side depending on the conditions of treatment. The dorsiventrality of the blade therefore influences but does not limit the direction of curvature. The dorsiventral directionality of response to growth regulators and the response to changes in the orientation to gravity are seen as indicating that blade curvatures are analogous to negative or positive gravitropism. It is noted that neither blade hyponasty or epinasty can be accounted for by ethylene alone. Petiole responses, however, are not directional, and the leaf angle changes induced by rotation or auxin treatment can be accounted for by ethylene production.

Hayes, A. B.↗

The Interaction of Calcium and Auxin in the Gravitropic Response of Roots

The role of calcium redistribution in the responding region of the root is examined, however, the potential connection between calcium and auxin redistribution in the elongation zone is not found. The following items are examined: (1) the effect of gravity on calcium movement across the elongation zone; (2) the effect of gravity on auxin movement across the elongation zone; and (3) the effect of calcium on auxin movement across the elongation zone. It is indicated that gravistimulation induces a physiological asymmetry in the auxin transport system of maize roots and that calcium increases the total transport of auxin across the root. Gravistimulation is apparently necessary for the enhancing effect of calcium on lateral auxin movement, and it is possible that the preferential downward movement of calcium across the elongation zone of gravistimulated roots plays a role in establishing the auxin asymmetry proposed to cause positive gravitropic curvature.

Evans, M. L.↗

Transduction of the Root Gravitropic Stimulus: Can Apical Calcium Regulate Auxin Distribution?

The hypothesis was tested that calcium, asymmetrically distributes in the root cap upon reorientation to gravity, affects auxin transport and thereby auxin distribution at the elongation zone. It is assumed that calcium exists in the root cap and is asymmetrically transported in root caps altered from a vertical to a horizontal position and that the meristem, the tissue immediately adjacent to the root cap and lying between the site of gravity perception and the site of gravity response, is essential for mediation of gravitropism. Tip calcium in root gravicurvature was implicated. The capstone evidence is that the root cap has the capacity to polarly translocate exogenous calcium downward when tissue is oriented horizontally, and that exogenous calcium, when supplied asymmetrically at the root tip, induces curvature and dictates the direction of curvature in both vertical and horizontal corn roots.

Edwards, K. L.↗

Gravitropism in Leafy Dicot Stems

A polarizing research microscope with rotating stage and associated camera equipment were ordered, and techniques of fixation and preparation of specimens were perfected for studying possible changes in orientation of cellulose microfibrils in cell walls of gravistimulated dicot stems. Acid ethephon solutions or acid without ethephon caused elongation of stem tissues where they were applied; stems bent away from the side of application. Acid solutions applied to the bottom of horizontal stems greatly delayed bending. Research in tissue sensitivity changes during gravitropic bending of soybean hypocotyls while immersed in auxin and in castor bean stems is also reported.

Salisbury, F. B.↗

Gravitropic responses of plants in the absence of a complicating G-force (6-IML-1)

On the Earth it is patently impossible to measure any tropistic, physiologic, or morphogenic reactions to environmental stimuli without taking into account our planet's gravitational influence on the time course of the test subject's response. It follows that all published reports of quantitative measurements of such responses must have been contaminated by an additional gravity dependent component which probably was not trivial. Our research effort has as its principal scientific objective, the acquisition of experimental data from tests in a microgravity environment that will address a number of basic questions about plants' gravitropic responses to the perception of transversely applied g forces in the hypogravity range, from essentially zero to unit g. Comparable tests on Earth but in the same flight hardware, referred to as the Gravitational Plant Physiology Facility (GPPF), will provide 1 g data for various useful comparisons. Four specific scientific questions are addressed.

Brown, Allan H.↗

A proposal to determine properties of the gravitropic response of plants in the absence of a complicating g-force (GTHRES)

Gravitropic responses of oat seedlings (Avena sativa L.) were measured on Earth and in microgravity (IML-1). The seedlings were grown at 1 g either on Earth or on 1 g centrifuges. They were challenged by centripetal accelerations for which the intensity and duration of the stimulations were varied. All stimulation intensities were in the hypogravity region from 0.1 to 1.0 g. All responses occurred either in Spacelab microgravity or during clinorotation on Earth. The experiments were carried out with the same apparatus in Spacelab and on Earth. The experiments addressed a series of scientific questions and useful data were obtained to provide answers to some but not all of those questions.

Brown, Allan H.↗