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At least 91 records · Page 5

Light regulation of the growth response in corn root gravitropism

Roots of Merit variety corn (Zea mays L.) require red light for orthogravitropic curvature. Experiments were undertaken to identify the step in the pathway from gravity perception to asymmetric growth on which light may act. Red light was effective in inducing gravitropism whether it was supplied concomitant with or as long as 30 minutes after the gravity stimulus (GS). The presentation time was the same whether the GS was supplied in red light or in darkness. Red light given before the GS slightly enhanced the rate of curvature but had little effect on the lag time or on the final curvature. This enhancement was expanded by a delay between the red light pulse and the GS. These results indicate that gravity perception and at least the initial transduction steps proceed in the dark. Light may regulate the final growth (motor) phase of gravitropism. The time required for full expression of the light enhancement of curvature is consistent with its involvement in some light-stimulated biosynthetic event.

NASA Discipline Number 40-50↗

Reduced gravitropic sensitivity in roots of a starch-deficient mutant of Nicotiana sylvestris

Gravitropism was studied in seedlings of Nicotiana sylvestris Speg. et Comes wild-type (WT) and mutant NS 458 which has a defective plastid phosphoglucomutase (EC 2.7.5.1.). Starch was greatly reduced in NS 458 compared to the WT, but small amounts of starch were detected in rootcap columella cells in NS 458 by light and electron microscopy. The roots of WT are more sensitive to gravity than mutant NS 458 roots since: (1) in mutant roots, curvature was reduced and delayed in the time course of curvature; (2) curvature of mutant roots was 24-56% that of WT roots over the range of induction periods tested; (3) in intermittent-stimulation experiments, curvature of mutant roots was 37% or less than that of WT roots in all treatments tested. The perception time, determined by intermittent-stimulation experiments, was < or = 5 s for WT roots and 30-60 s for mutant roots. The growth rates for WT and NS 458 roots were essentially equal. These results and our previous results with WT and starchless mutant Arabidopsis roots (Kiss et al. 1989, Planta 177, 198-206) support the conclusions that a full complement of starch is necessary for full gravitropic sensitivity and that amyloplasts function in gravity perception. Since a presumed relatively small increase in plastid buoyant mass (N. sylvestris mutant versus Arabidopsis mutant) significantly improves the orientation of the N. sylvestris mutant roots, we suggest that plastids are the likeliest candidates to be triggering gravity perception in roots of both mutants.

Non-NASA Center↗

Phytochromes play a role in phototropism and gravitropism in Arabidopsis roots

Phototropism as well as gravitropism plays a role in the oriented growth of roots in flowering plants. In blue or white light, roots exhibit negative phototropism, but red light induces positive phototropism in Arabidopsis roots. Phytochrome A (phyA) and phyB mediate the positive red-light-based photoresponse in roots since single mutants (and the double phyAB mutant) were severely impaired in this response. In blue-light-based negative phototropism, phyA and phyAB (but not phyB) were inhibited in the response relative to the WT. In root gravitropism, phyB and phyAB (but not phyA) were inhibited in the response compared to the WT. The differences observed in tropistic responses were not due to growth limitations since the growth rates among all the mutants tested were not significantly different from that of the WT. Thus, our study shows that the blue-light and red-light systems interact in roots and that phytochrome plays a key role in plant development by integrating multiple environmental stimuli. c2003 COSPAR. Published by Elsevier Ltd. All rights reserved.

Non-NASA Center↗

Geoperception in primary and lateral roots of Phaseolus vulgaris (Fabaceae). III. A model to explain the differential georesponsiveness of primary and lateral roots

Half-tipped primary and lateral roots of Phaseolus vulgaris bend toward the side of the root on which the intact half tip remains. Therefore, tips of lateral and primary roots produce growth effectors capable of inducing gravicurvature. The asymmetrical placement of a tip of a lateral root onto a detipped primary root results in the root bending toward the side of the root onto which the tip was placed. That is, the lesser graviresponsiveness of lateral roots as compared with primary roots is not due to the inability of their caps to produce growth inhibitors. The more pronounced graviresponsiveness of primary roots is positively correlated with the presence of columella tissues that are 3.8 times longer, 1.7 times wider, and 10.5 times more voluminous than the columellas of lateral roots. We propose that the lack of graviresponsiveness exhibited by lateral roots is due to the fact that they (i) produce smaller amounts of the inhibitor than primary (i.e., strongly graviresponsive) roots and (ii) are unable to redistribute the inhibitor so as to be able to create a concentration gradient sufficient to induce a pronounced gravitropic response.

NASA Program Space Biology↗

The effect of the external medium on the gravitropic curvature of rice (Oryza sativa, Poaceae) roots

The roots of rice seedlings, growing in artificial pond water, exhibit robust gravitropic curvature when placed perpendicular to the vector of gravity. To determine whether the statolith theory (in which intracellular sedimenting particles are responsible for gravity sensing) or the gravitational pressure theory (in which the entire protoplast acts as the gravity sensor) best accounts for gravity sensing in rice roots, we changed the physical properties of the external medium with impermeant solutes and examined the effect on gravitropism. As the density of the external medium is increased, the rate of gravitropic curvature decreases. The decrease in the rate of gravicurvature cannot be attributed to an inhibition of growth, since rice roots grown in 100 Osm/m3 (0.248 MPa) solutions of different densities all support the same root growth rate but inhibit gravicurvature increasingly with increasing density. By contrast, the sedimentation rate of amyloplasts in the columella cells is unaffected by the external density. These results are consistent with the gravitational pressure theory of gravity sensing, but cannot be explained by the statolith theory.

Non-NASA Center↗

Tomato growth as affected by root-zone temperature and the addition of gibberellic acid and kinetin to nutrient solutions

The effect of root-zone temperature on young tomato plants (Lycopersicon esculentum Mill. cv. Heinz 1350) was evaluated in controlled environments using a recirculating solution culture system. Growth rates were measured at root-zone temperatures of 15 degrees, 20 degrees, 25 degrees, and 30 degrees C in a near optimum foliar environment. Optimum growth occurred at 25 degrees to 30 degrees during the first 4 weeks of growth and 20 degrees to 25 degrees during the 5th and 6th weeks. Growth was severely restricted at 15 degrees. Four concentrations of gibberellic acid (GA3) and kinetin were added to the nutrient solution in a separate trial; root-zone temperature was maintained at 15 degrees and 25 degrees. Addition of 15 micromoles GA3 to solutions increased specific leaf area, total leaf area, and dry weight production of plants in both temperature treatments. GA3-induced growth stimulation was greater at 15 degrees than at 25 degrees. GA3 may promote growth by increasing leaf area, enhancing photosynthesis per unit leaf area, or both. Kinetic was not useful in promoting growth at either temperature.

NASA Program CELSS↗

Patterns of auxin and abscisic acid movement in the tips of gravistimulated primary roots of maize

Because both abscisic acid (ABA) and auxin (IAA) have been suggested as possible chemical mediators of differential growth during root gravitropism, we compared with redistribution of label from applied 3H-IAA and 3H-ABA during maize root gravitropism and examined the relative basipetal movement of 3H-IAA and 3H-ABA applied to the caps of vertical roots. Lateral movement of 3H-ABA across the tips of vertical roots was non-polar and about 2-fold greater than lateral movement of 3H-IAA (also non-polar). The greater movement of ABA was not due to enhanced uptake since the uptake of 3H-IAA was greater than that of 3H-ABA. Basipetal movement of label from 3H-IAA or 3H-ABA applied to the root cap was determined by measuring radioactivity in successive 1 mm sections behind the tip 90 minutes after application. ABA remained largely in the first mm (point of application) whereas IAA was concentrated in the region 2-4 mm from the tip with substantial levels found 7-8 mm from the tip. Pretreatment with inhibitors of polar auxin transport decreased both gravicurvature and the basipetal movement of IAA. When roots were placed horizontally, the movement of 3H-IAA from top to bottom across the cap was enhanced relative to movement from bottom to top whereas the pattern of movement of label from 3H-ABA was unaffected. These results are consistent with the hypothesis that IAA plays a role in root gravitropism but contrary to the idea that gravi-induced asymmetric distribution of ABA contributes to the response.

Non-NASA Center↗

Growth and microtubule orientation of Zea mays roots subjected to osmotic stress

Previous work has shown that microtubule (MT) reorientation follows the onset of growth inhibition on the lower side of graviresponding roots, indicating that growth reduction can occur independently of MT reorientation. To test this observation further, we examined whether the reduction in growth in response to osmotic stress is correlated with MT reorientation. The distribution and rate of growth in maize roots exposed to 350 mOsm sorbitol and KCl or 5 mM Mes/Tris buffer were measured with a digitizer. After various times roots were processed for indirect immunofluorescence microscopy. Application of sorbitol or KCl had no effect on the organization of MTs in the apical 2 mm of the root but resulted in striking and different effects in the basal region of the root. Sorbitol treatment caused rapid appearance of oval to circular holes in the microtubular array that persisted for at least 9 h. Between 30 min and 4 h of submersion in KCl, MTs in cortical cells 4 mm and farther from the quiescent center began to reorient oblique to the longitudinal axis. After 9 h, the alignment of MTs had shifted to parallel to the root axis but MTs of the epidermal cells remained transverse. In KCl-treated roots MT reorientation appeared to follow a pattern of development similar to that in controls but without elongation. Our data provide additional evidence that MT reorientation is not the cause but a consequence of growth inhibition.

NASA Discipline Plant Biology↗

Root gravitropism: an experimental tool to investigate basic cellular and molecular processes underlying mechanosensing and signal transmission in plants

The ability of plant organs to use gravity as a guide for growth, named gravitropism, has been recognized for over two centuries. This growth response to the environment contributes significantly to the upward growth of shoots and the downward growth of roots commonly observed throughout the plant kingdom. Root gravitropism has received a great deal of attention because there is a physical separation between the primary site for gravity sensing, located in the root cap, and the site of differential growth response, located in the elongation zones (EZs). Hence, this system allows identification and characterization of different phases of gravitropism, including gravity perception, signal transduction, signal transmission, and curvature response. Recent studies support some aspects of an old model for gravity sensing, which postulates that root-cap columellar amyloplasts constitute the susceptors for gravity perception. Such studies have also allowed the identification of several molecules that appear to function as second messengers in gravity signal transduction and of potential signal transducers. Auxin has been implicated as a probable component of the signal that carries the gravitropic information between the gravity-sensing cap and the gravity-responding EZs. This has allowed the identification and characterization of important molecular processes underlying auxin transport and response in plants. New molecular models can be elaborated to explain how the gravity signal transduction pathway might regulate the polarity of auxin transport in roots. Further studies are required to test these models, as well as to study the molecular mechanisms underlying a poorly characterized phase of gravitropism that is independent of an auxin gradient.

Non-NASA Center↗

Flexible Microsensor Array for the Root Zone Monitoring of Porous Tube Plant Growth System

Control of oxygen and water in the root zone is vital to support plant growth in the microgravity environment. The ability to control these sometimes opposing parameters in the root zone is dependent upon the availability of sensors to detect these elements and provide feedback for control systems. In the present study we demonstrate the feasibility of using microsensor arrays on a flexible substrate for dissolved oxygen detection, and a 4-point impedance microprobe for surface wetness detection on the surface of a porous tube (PT) nutrient delivery system. The oxygen microsensor reported surface oxygen concentrations that correlated with the oxygen concentrations of the solution inside the PT when operated at positive pressures. At negative pressures the microsensor shows convergence to zero saturation (2.2 micro mol/L) values due to inadequate water film formation on porous tube surface. The 4-point microprobe is useful as a wetness detector as it provides a clear differentiation between dry and wet surfaces. The unique features of the dissolved oxygen microsensor array and 4-point microprobe include small and simple design, flexibility and multipoint sensing. The demonstrated technology is anticipated to provide low cost, and highly reliable sensor feedback monitoring plant growth nutrient delivery system in both terrestrial and microgravity environments.

Sathyan, Sandeep↗

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↗

Analysis of growth patterns during gravitropic curvature in roots of Zea mays by use of a computer-based video digitizer

A computer-based video digitizer system is described which allows automated tracking of markers placed on a plant surface. The system uses customized software to calculate relative growth rates at selected positions along the plant surface and to determine rates of gravitropic curvature based on the changing pattern of distribution of the surface markers. The system was used to study the time course of gravitropic curvature and changes in relative growth rate along the upper and lower surface of horizontally-oriented roots of maize (Zea mays L.). The growing region of the root was found to extend from about 1 mm behind the tip to approximately 6 mm behind the tip. In vertically-oriented roots the relative growth rate was maximal at about 2.5 mm behind the tip and declined smoothly on either side of the maximum. Curvature was initiated approximately 30 min after horizontal orientation with maximal (50 degrees) curvature being attained in 3 h. Analysis of surface extension patterns during the response indicated that curvature results from a reduction in growth rate along both the upper and lower surfaces with stronger reduction along the lower surface.

NASA Discipline Plant Biology↗

Evidence for a role in growth and salt resistance of a plasma membrane H+-ATPase in the root endodermis

The plasma membrane of plant cells is energized by an electrochemical gradient produced by P-type H+-ATPases (proton pumps). These pumps are encoded by at least 12 genes in Arabidopsis. Here we provide evidence that isoform AHA4 contributes to solute transport through the root endodermis. AHA4 is expressed most strongly in the root endodermis and flowers, as suggested by promoter-GUS reporter assays. A disruption of this pump (aha4-1) was identified as a T-DNA insertion in the middle of the gene (after VFP(574)). Truncated aha4-1 transcripts accumulate to approximately 50% of the level observed for AHA4 mRNA in wild-type plants. Plants homozygous for aha4-1 (-/-) show a subtle reduction in root and shoot growth compared with wild-type plants when grown under normal conditions. However, a mutant phenotype is very clear in plants grown under salt stress (e.g., 75 or 110 mM NaCl). In leaves of mutant plants subjected to Na stress, the ratio of Na to K increased 4-5-fold. Interestingly, the aha4-1 mutation appears to be semidominant and was only partially complemented by the introduction of additional wild-type copies of AHA4. These results are consistent with the hypothesis that aha4-1 may produce a dominant negative protein or RNA that partially disrupts the activity of other pumps or functions in the root endodermal tissue, thereby compromising the function of this cell layer in controlling ion homeostasis and nutrient transport.

NASA Discipline Plant Biology↗

A growth analysis of waterlogging damage in mung bean (Phaseolus aureus)

Mung beans (Phaseolus aureus Roxb.) were grown for 2 weeks in gravel-vermiculite soilless mix in a growth chamber and subjected to a 1-week waterlogging period followed by a 1-week recovery period. Sequential harvests were made to determine the time course of effects of waterlogging and subsequent recovery on growth parameters by techniques of growth analysis. Root dry matter was the first to be affected, along with an increase in leaf dry matter and specific leaf weight. After a 1-week waterlogging period, specific leaf weight had more than doubled in the stressed plants. Leaf area declined in relation to the control plants as did the ratio of root dry matter to shoot dry matter. During the recovery period there was an increase in the dry matter allocation to the roots relative to the shoot. Specific leaf weight fell to control levels although the rate of leaf area elaboration did not increase during this time, suggesting a redistribution of stored assimilates from the leaves. Net assimilation rate increased during the waterlogging period, probably due to a restriction in root metabolism and reduced translocation out of the leaf rather than to an increase in photosynthesis. Net assimilation rate of waterlogged plants was severely reduced compared with control plants during the recovery period. Both relative growth rate and leaf area duration declined during the waterlogging period and declined further subsequent to the waterlogging treatment. The results illustrate the interrelationships between root and shoot carbon budgets in mung bean during response to the stress of waterlogging.

Non-NASA Center↗

Phytochromes A and B mediate red-light-induced positive phototropism in roots

The interaction of tropisms is important in determining the final growth form of the plant body. In roots, gravitropism is the predominant tropistic response, but phototropism also plays a role in the oriented growth of roots in flowering plants. In blue or white light, roots exhibit negative phototropism that is mediated by the phototropin family of photoreceptors. In contrast, red light induces a positive phototropism in Arabidopsis roots. Because this red-light-induced response is weak relative to both gravitropism and negative phototropism, we used a novel device to study phototropism without the complications of a counteracting gravitational stimulus. This device is based on a computer-controlled system using real-time image analysis of root growth and a feedback-regulated rotatable stage. Our data show that this system is useful to study root phototropism in response to red light, because in wild-type roots, the maximal curvature detected with this apparatus is 30 degrees to 40 degrees, compared with 5 degrees to 10 degrees without the feedback system. In positive root phototropism, sensing of red light occurs in the root itself and is not dependent on shoot-derived signals resulting from light perception. Phytochrome (Phy)A and phyB were severely impaired in red-light-induced phototropism, whereas the phyD and phyE mutants were normal in this response. Thus, PHYA and PHYB play a key role in mediating red-light-dependent positive phototropism in roots. Although phytochrome has been shown to mediate phototropism in some lower plant groups, this is one of the few reports indicating a phytochrome-dependent phototropism in flowering plants.

NASA Discipline Plant Biology↗

Protein synthesis in geostimulated root caps

A study is presented of the processes occurring in the root cap of corn which are requisite for the formation of root cap inhibitor and which can be triggered or modulated by both light and gravity. The results of this study indicate the importance of protein synthesis for light-induced gravitropic bending in roots. Root caps in which protein synthesis is prevented are unable to induce downward bending. This suggests that light acts by stimulating proteins which are necessary for the translation of the gravitropic stimulus into a growth response (downward bending). The turnover of protein with time was also examined in order to determine whether light acts by stimulating the synthesis of unique proteins required for downward growth. It is found that auxin in combination with light allows for the translation of the gravitropic stimulus into a growth response at least in part through the modification of protein synthesis. It is concluded that unique proteins are stimulated by light and are involved in promoting the downward growth in roots which are responding to gravity.

Feldman, L. J.↗

The role of calcium and calmodulin in the response of roots to gravity

There is general agreement that, in roots, the primary detection of the gravitropic signal occurs in the columella cells of the cap and that this results in the generation of a signal that moves into the elongation zone causing the asymmetric growth that leads to downward curvature. Recent work has generated considerable evidence that indicates that auxin is the ultimate mediator of differential growth during root (and shoot) gravitropism. Our studies of the time course of curvature, auxin redistribution and/or adaptation, and electrical potential changes in maize roots have led to the following generalizations: (1) downward curvature begins 18 to 32 min following gravistimulation; (2) asymmetric auxin redistribution across the root cap begins at about the same time as curvature or perhaps slight earlier; (3) there is a lag of approx. 15 min in the response of roots to applied auxin; and (4) gravi-induced changes in intracellular potentials of cortical cells within the elongation zone occur within 30 s following stimulation.

Evans, Michael L.↗