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PubMed1972

Migration of Calcium and Its Role in the Regulation of Seismonasty in the Motor Cell of Mimosa pudica L.

Volume and conformational changes of the contractile tannin vacuoles of the abaxial motor cells of the primary pulvinus of Mimosa pudica L. parallel the seismonastic leaf movement. Since such changes in cells and organelles of animal systems are often regulated by calcium, we studied Ca(2+) movement in the motor cells and tissue. By fixation with Lillie's neutral buffered formalin, followed by staining with alizarin red sulfate (ARS), calcium was localized in the tannin vacuoles of the motor cells of the primary pulvinus. After treatment with ethylenediaminetetraacetate, 8-hydroxyquinoline, and several other calcium-complexing or extracting agents, the color reaction due to alizarin red sulfonate was no longer present. By using an analytical method, it was shown that the effluent from stimulated pulvini has significantly more Ca(2+) than that from unstimulated controls. Ten millimolar LaCl(3) inhibits recovery of the tannin vacuole in vivo in 10 mm CaCl(2) or in distilled water. Quantitative data obtained by microspectrophotometry demonstrated calcium migration during the bending movement of the primary pulvinus. In the adaxial motor cells a small amount of calcium migrates from the tannin vacuole, and calcium on the cell wall moves to the central vacuole. In the abaxial half, a large amount of calcium from the tannin vacuole moves to the central vacuole of the motor cell. It is probable that the calcium binds to the microfibrillar contents of the central vacuole. These observations support the contention that Ca(2+) migrates between the surface of the tannin vacuole and the inside of the central vacuole. The recovery and maintenance of the tannin vacuole in the spherical form may play a role in maintaining turgor in the motor cells of the abaxial half of the primary pulvinus of Mimosa.

Journal Article↗
PubMed1993

Molecular basis of the increase in invertase activity elicited by gravistimulation of oat-shoot pulvini.

An asymmetric (top vs. bottom) increase in invertase activity is elicited by gravistimulation in oat-shoot pulvini starting within 3 h after treatment. In order to analyze the regulation of invertase gene expression in this system, we examined the effect of gravistimulation on invertase mRNA induction. Total RNA and poly (A)+RNA, isolated from oat pulvini, and two oligonucleotide primers, corresponding to two conserved amino-acid sequences (NDPNG and WECPD) found in invertase from other species, were used for the polymerase chain reaction (PCR). A partial-length cDNA (550 base pairs) was obtained and characterized. There was a 52% deduced amino-acid sequence homology to that of carrot beta-fructosidase and a 48% homology to that of tomato invertase. Northern blot analysis showed that there was an obvious transient accumulation of invertase mRNA elicited by gravistimulation of oat pulvini. The mRNA was rapidly induced to a maximum level at 1 h following gravistimulation treatment and gradually decreased afterwards. The mRNA level in the bottom half of the oat pulvinus was significantly higher (five-fold) than that in the top half of the pulvinus tissue. The induction of invertase mRNA was consistent with the transient enhancement of invertase activity during the graviresponse of the pulvinus. These data indicate that the expression of the invertase gene(s) could be regulated by gravistimulation at the transcriptional and/or translational levels. Southern blot analysis showed that there were four genomic DNA fragments hybridized to the invertase cDNA. This suggests that an invertase gene family may exist in oat plants.

Amino Acid Sequence↗
PubMed2004

Biochemical and immunohistochemical characterization of Mimosa annexin.

To characterize the biochemical properties of plant annexin, we isolated annexin from Mimosa pudica L. and analyzed the biochemical properties conserved between Mimosa annexin and animal annexins, e.g. the ability to bind phospholipid and F-actin in the presence of calcium. We show that Mimosa annexin is distributed in a wide variety of tissues. Immunoblot analysis also revealed that the amount of annexin is developmentally regulated. To identify novel functions of Mimosa annexin, we examined the pattern of distribution and the regulation of its expression in the pulvinus. The amount of annexin in the pulvinus increased at night and was sensitive to abscisic acid; however, there was no detectable induction of annexin by cold or mechanical stimulus. Annexin distribution in the cell periphery during the daytime was changed to a cytoplasmic distribution at night, indicating that Mimosa annexin may contribute to the nyctinastic movement in the pulvinus.

Amino Acid Sequence↗
PubMed1982

Elemental analysis of freeze-dried thin sections of Samanea motor organs: barriers to ion diffusion through the apoplast.

Leaflet movements in the legume Samanea saman are dependent upon massive redistribution of potassium (K), chloride (Cl), and other solutes between opposing (extensor and flexor) halves of the motor organ (pulvinus). Solutes are known to diffuse through the apoplast during redistribution. To test the possibility that solute diffusion might be restricted by apoplastic barriers, we analyzed elements in the apoplast in freeze-dried cryosections of pulvini using scanning electron microscopy/x-ray microanalysis. Large discontinuities in apoplastic K and Cl at the extensor-flexor interface provide evidence for a barrier to solute diffusion. The barrier extends from the epidermis on upper and lower sides of the pulvinus to cambial cells in the central vascular core. It is completed by hydrophobic regions between phloem and cambium, and between xylem rays and surrounding vascular tissue, as deduced by discontinuities in apoplastic solutes and by staining of fresh sections with lipid-soluble Sudan dyes. Thus, symplastic pathways are necessary for ion redistribution in the Samanea pulvinus during leaflet movement. In pulvini from leaflets in the closed state, all cells on the flexor side of the barrier have high internal as well as external K and Cl, whereas cells on the extensor side have barely detectable internal or external K or Cl. Approximately 60% of these ions are known to migrate to the extensor during opening; all return to the flexor during subsequent closure. We propose that solutes lost from shrinking cells in the outer cortex diffuse through the apoplast to plasmodesmata-rich cells of the inner cortex, collenchyma, and phloem; and that solutes cross the barrier by moving through plasmodesmata.

Cell Wall↗
PubMed1988

Movement of water in conjunction with plant movement visualized by NMR imaging.

The water distribution in the pulvinus of Mimosa can be visualized by an NMR imaging technique. After stimulation of a Mimosa plant, water in the lower half of the main pulvinus disappeared, the water previously contained in this area seeming to be transferred to the upper half of the main pulvinus. Movement of the water in conjunction with Mimosa movement was visualized sequentially by a non-invasive NMR imaging procedure.

Magnetic Resonance Spectroscopy↗
PubMed1978

Phototropic Response to Vectorial Light in Leaves of Lavatera cretica L.

The mechanism by which the mature leaf of certain plants reorients its lamina to face the sun throughout the day was studied in Lavatera cretica L. The photoreceptor for this response differs fundamentally from the one involved in the phototropic growth response, by sensing light as a vector, rather than as a difference in luminous flux. The photoreceptor is located in the veins, which radiate in the plane of the lamina from the pulvinus situated at the junction between the lamina and petiole. The integrated response to the messages from the different veins takes place by differential turgor changes in a motor tissue surrounding the central vascular cylinder of the pulvinus, in which the veins coalesce. The differential turgor in the different segments of the motor tissue determines the orientation of the lamina. The photoreceptor reacts only to a parallel light beam striking the vein obliquely (from above). When half of the lamina is shaded, the leaf does not reorient in response to perpendicular illumination and its reorientation in response to an oblique beam is slower and partial, to a greater extend when the half-leaf is centrifugally illuminated than when it is centripetally illuminated. Application of 2,3,5 tri-iodobenzoic acid to the base of the veins in the shaded half-leaf eliminated all restrictions from the response to centrifugal illumination and totally inhibited the response to centripetal illumination. The results are consistent with a hypothesis that centrifugally illuminated veins generate turgor in their associated motor tissue in the pulvinus by activating K(+) uptake, while centripetally illuminated veins cause loss of turgor in their associated motor tissue by deactivating K(+) uptake, which leads to passive leakage of K(+). When the entire lamina is exposed to oblique illumination, the centrifugally illuminated half and the centripetally illuminated half cooperate in the full response. Shaded parts of the lamina apparently interfere with the response by supplying their associated motor tissue with auxin, which presumably causes in it an active export of protons and concomitant uptake of K(+), thereby establishing a static "dark turgor" in it.

Journal Article↗
PubMed1981

Changes in Endogenous Gibberellins and the Metabolism of [H]GA(4) after Geostimulation in Shoots of the Oat Plant (Avena sativa).

The recovery from "lodging," or bending over, by shoots of 42-day-old Avena sativa plants is controlled primarily by a negatively geotropic differential growth of the lower halves of the p-1 node-pulvinus and the base of the p-1 internode, relative to the upper halves. Although geostimulation causes a significant reduction in p-1 internode length, dry matter accumulation in the p-1 node-pulvinus is increased, apparently at the expense of the sheath. Recovery to an angle of 30 degrees is associated with changes in endogenous gibberellin-like substances (GAs), and in differential metabolism of applied [(3)H]GA(4) (1.4 Curie per millimole). Although geostimulation depressed total GAs (relative to upright plant parts) to 0.40 and 0.13 for node-pulvini and sheaths, respectively, it increased them 2-fold for internodes. Within the plant part geostimulation increased GAs (relative to upper halves) 29- and 7-fold in lower halves of node-pulvini and internodes, respectively, but reduced GAs to 0.3 in lower halves of sheaths. At age 42 days a GA(4/7)-like (nonpolar) substance predominates, with lesser amounts of a GA(3)-like (polar) substance. Native GAs of Avena include GA(3), GA(4), and GA(7). Geostimulation enhanced the ratio of nonpolar to polar GAs for both halves of internodes, but tended to depress it for sheaths and nodepulvini.The disposition and metabolism of applied [(3)H]GA(4) confirmed the above trends for endogenous GAs regarding localization (e.g. up to 2-fold increases in [(3)H]GA(4) and acidic (3)H-metabolites in the lower halves, relative to upper halves). Also, metabolism into highly water-soluble (3)H-metabolites (biologically inactive conjugates?) was greater (up to 1.8-fold) in upper than in lower halves. The end result of such metabolic trends would be to reduce acidic (biologically active?) GAs in the upper half, while retaining them for a longer time in the lower half.Geotropically stimulated Avena shoots thus increase, within 24 hours, the levels of acidic GAs in the lower halves of the p-1 node-pulvinus and p-1 internode, the two plant parts responsible for the geostimulated growth.

Journal Article↗
PubMed2001

Blue-light-dependent osmoregulation in protoplasts of Phaseolus vulgaris Pulvini.

Blue light was found to induce shrinkage of the protoplasts isolated from first-leaf lamina pulvini of 18-day-old Phaseolus vulgaris. The response was transient following pulse stimulation, while it was sustainable during continuous stimulation. No apparent difference was found between flexor and extensor protoplasts. Protoplasts of the petiolar segment located close to the pulvinus showed no detectable response. In the plants used, the pulvinus was fully matured and the petiole was ceasing its elongation growth. When younger, 12-day-old, plants were used, however, the petiolar protoplasts did respond to blue light. The pulse-induced response was similar to that in pulvinar protoplasts, although the response to continuous stimulation was transient and differed from that in pulvinar protoplasts. No shrinkage was induced in pulvinar protoplasts when the far-red-light-absorbing form of phytochrome was absent for a period before blue-light stimulation, indicating that the blue-light responsiveness is strictly controlled by phytochrome. Inhibitors of anion channels and H(+)-ATPase abolished the shrinking response, supporting the view that protoplasts shrink by extruding ions. The response of pulvinar protoplasts is probably involved in the blue-light-induced, turgor-based movement of pulvini. The blue-light responding system in pulvini is suggested to have evolved from that functioning in other growing organs.

Adaptation, Physiological↗
PubMed2006

Change in the actin cytoskeleton during seismonastic movement of Mimosa pudica.

The seismonastic movement of Mimosa pudica is triggered by a sudden loss of turgor pressure. In the present study, we compared the cell cytoskeleton by immunofluorescence analysis before and after movement, and the effects of actin- and microtubule-targeted drugs were examined by injecting them into the cut pulvinus. We found that fragmentation of actin filaments and microtubules occurs during bending, although the actin cytoskeleton, but not the microtubules, was involved in regulation of the movement. Transmission electron microscopy revealed that actin cables became loose after the bending. We injected phosphatase inhibitors into the severed pulvinus to examine the effects of such inhibitors on the actin cytoskeleton. We found that changes in actin isoforms, fragmentation of actin filaments and the bending movement were all inhibited after injection of a tyrosine phosphatase inhibitor. We thus propose that the phosphorylation status of actin at tyrosine residues affects the dynamic reorganization of actin filaments and causes seismonastic movement.

Actins↗
PubMed1988

Effect of dark pretreatment on the kinetics of response of barley pulvini to gravistimulation and hormones.

Starch in pulvinus amyloplasts of barley (Hordeum vulgare cv Larker) disappears when 45-day-old, light-grown plants are given 5 days of continuous darkness. The effect of this loss on the pulvinus graviresponse was evaluated by following changes in the kinetics of response during the 5-day dark period. Over 5 days of dark pretreatment, the lag to initial graviresponse and the subsequent half-time to maximum steady state bending rate increased significantly while the maximum bending rate did not change. The change in response to applied indoleacetic acid (100 micromolar) plus gibberellic acid (10 micromolar) without gravistimulation, under identical dark pretreatments, was used as a model system for the response component of gravitropism. Dark pretreatment did not change the lag to initial response following hormone application to vertical pulvini, but both the maximum bending rate and the half-time to the maximum rate were significantly reduced. Also, after dark pretreatment, significant bending responses following hormone application were observed in vertical segments with or without added sucrose, while gravistimulation produced a response only if segments were given sucrose. These results indicate that starch-filled amyloplasts are required for the graviresponse of barley pulvini and suggest that they function in the stimulus perception and signal transduction components of gravitropism.

Darkness↗
PubMed1981

Apoplastic transport of ions in the motor organ of Samanea.

Turgor-mediated leaf movements of the legume Samanea saman are associated with the migration of K(+) and Cl(-) between opposing sides of the motor organs (pulvini). We have investigated the pathway of this ion migration by localizing K(+) and Cl(-) within the secondary pulvinus at various times during leaf movements. Ion distributions in freeze-dried cryosections of pulvini were determined by scanning electron microscopy/x-ray microanalysis. The results indicate that the apoplast is a major pathway for the migration of K(+) and Cl(-) within the pulvinus.

Journal Article↗
PubMed2006

Phosphorylation of SPICK2, an AKT2 channel homologue from Samanea motor cells.

SPICK2, a homologue of the weakly-inward-rectifying Shaker-like Arabidopsis K channel, AKT2, is a candidate K+-influx channel participating in light- and clock-regulated leaf movements of the legume, Samanea saman. Light and the biological clock regulate the in situ K+-influx channel activity differentially in extensor and flexor halves of the pulvinus (the S. saman leaf motor organ), and also-though differently-the transcript level of SPICK2 in the pulvinus. This disparity between the in situ channel activity versus its candidate transcript, along with the sequence analysis of SPICK2, suggest an in situ regulation of the activity of SPICK2, possibly by phosphorylation and/or by interaction with cAMP. Consistent with this (i) the activity of the voltage-dependent K+-selective fraction of the inward current in extensor and flexor cells was affected differentially in whole-cell patch-clamp assays promoting phosphorylation (using the protein phosphatase inhibitor okadaic acid); (ii) several proteins in isolated plasma membrane-enriched vesicles of the motor cells underwent phosphorylation without an added kinase in conditions similar to patch-clamp; and (iii) the SPICK2 protein was phosphorylated in vitro by the catalytic subunit of the broad-range cAMP-dependent protein kinase. All of these results are consistent with the notion that SPICK2 is the K+-influx channel, and is regulated in vivo directly by phosphorylation.

Animals↗
PubMed1994

A Role for the Stele in Intertissue Signaling in the Initiation of Abscission in Bean Leaves (Phaseolus vulgaris L.).

A combination of microdissection and viscometric endo-[beta]-1,4-glucanhydrolase assays was used to investigate if the early appearance of the abscission-related isoelectric point-9.5 endo-[beta]-1,4-glucanhydrolase in the stele of the pulvinus and abscission zone of the foliar abscission zone of Phaseolus vulgaris L. prior to cell separation (reported by E. del Campillo, P.D. Reid, R. Sexton, L.N.Lewis [1990] Plant Cell 2: 245-254) indicates that the vascular tissue of this region has a specific role in abscission. We find that no endo-[beta]-1,4-glucanhydrolase activity or cell separation is detectable in the abscission zone cortex if the abscission zone cortex is separated from the stele tissue. If the stele is separated from the abscission zone cortex after a lag period but again before any endo-[beta]-1,4-glucanhydrolase activity is present in the abscission zone cortex, then the enzyme is produced in the cortex and abscission ensues. We conclude that the cortex of the abscission zone is able to abscind independently of the vascular tissue only after the vascular tissue has begun to respond to abscission-promoting signals. We suggest that ethylene promotes formation of an abscission-permitting signal in the stele of the abscission zone and pulvinus, and that this signal is an essential elicitor for the synthesis of cell separation enzymes in the target cells of the abscission zone cortex.

Journal Article↗
PubMed1976

Studies on acidification of media by Avena stem segments in the presence and absence of gibberellic Acid.

The rate of acidification of media by Avena stem segments was studied with a titrimeter. GA(3) increased this rate by an average of 17% if supplied to the segments 90 min prior to measurement. GA(3) inhibited the rate by 15% if supplied 10 min prior to measurement. After 90 min incubation, stimulation of elongation had started; at 10 min, GA(3) had not yet started to stimulate elongation in the segments.The acidification rates of the nodes (including the sheath-pulvinus), leaf sheath bases, and the internode bases of the stem segments were determined for plus and minus GA(3)-treated segments. The internode fraction contributes most to modification of the acidification rate, the node-pulvinus fraction less so, and the nongrowing sheath not at all.Acidification rates were measured for segments in different stages of elongation (lag, log, and plateau phases of growth). Segments in these growth stages were obtained from intact plants and from segments preincubated in sucrose and sucrose + GA(3). Segments from all sources which are in the log phase of growth have the highest rates, those in the plateau phase the lowest. For lag and log growth phases, segments preincubated in sucrose + GA(3) show the highest rates, those preincubated in sucrose the lowest rates. The opposite occurs for segments in the plateau phase of growth.Segments stimulated to grow by GA(3) cause the pH of their incubation media to drop to pH 5.15 from an initial pH of 6.5. Nonstimulated segments cause a drop to pH 5.6. Long term growth of the segments is maximal in media buffered to pH 5 in the presence and absence of GA(3).Our results support the idea that GA(3) stimulates an active acidification process in Avena stem segments just after GA(3) starts to stimulate growth in the segments, and that such an acidification process could play an important role in wall-loosening during active growth of the internode.

Journal Article↗
PubMed1979

Fatty Acids and Circadian Rhythms in Phaseolus coccineus: Effects of Light, Temperature, and Chemicals.

Five major fatty acids, palmitic (16:0), stearic (18:0), oleic (18:1), linoleic (18:2), and linolenic (18:3), were identified in polar lipid extracts from pulvini of Samanea saman and Phaseolus coccineus. In P. coccineus their distribution varied quantitatively in the laminar pulvinus, petiolar pulvinus, petiole, stem, leaf and root. Short pulses of red light did not greatly affect the relative quantities of fatty acids in dark grown P. coccineus, but a 30-minute exposure of red light generally increased the degree of unsaturation by increasing linolenic acid and decreasing linoleic and palmitic acids.P. coccineus seeds were exposed to several substituted pyridazinones as well as cerulenin and dimethylethanolamine. The pyridazinones San 6706 and norflurazon altered fatty acid composition but also altered morphology and inhibited chlorophyll synthesis. Exposure to 10 C for 72 hours caused a small but significant increase in the degree of unsaturation of P. coccineus fatty acids but results were equivocal with S. saman.

Journal Article↗
PubMed1984

Extensor and flexor protoplasts from samanea pulvini : I. Isolation and initial characterization.

Protoplasts were isolated from extensor and flexor regions of open pulvini of the nyctinastic tree Samanea saman. Both types of protoplasts undergo many changes during isolation. Extensor protoplasts are univacuolate in vivo, but some become multivacuolate. All flexor protoplasts are univacuolate. In an open pulvinus, extensor cells have a higher osmotic pressure than flexor cells. However, both types of protoplasts can be isolated with optimal yield using the same osmoticum (0.5 molar sorbitol) in the digestion medium. This suggests that some leakage of osmoticum occurs during harvest or digestion, especially from extensor tissue. Despite these changes, both types of protoplasts extrude protons in response to 10 micromolar fusicoccin (1.6-1.8 nanoequivalent/10(6) protoplasts/minute), demonstrating that the protoplasts are metabolically active and that proton transport mechanisms must be at least partially functional. The changes in vacuolar structure and osmotic pressure are what one might expect if the protoplasts, which are isolated from open pulvini, take on characteristics of cells in a closed pulvinus.

Journal Article↗
PubMed1987

Water Relations in Pulvini from Samanea saman: I. Intact Pulvini.

The movement of Samanea leaflets depends upon changes in the curvature of the pulvinus at the base of each leaflet. Pulvinar bending and straightening, in turn, are driven by the movement of water between opposing (extensor and flexor) sides of the pulvinus. Although water movement depends on water potential (Psi) and thus on osmotic potential (pi) and hydrostatic pressure (P), none of these parameters have been measured in Samanea. In this investigation, Psi and pi were measured and P was calculated for extensor and flexor tissues of excised, whole pulvini that were open in the light and closed in the dark. In fully open pulvini, pi in the extensor was generally between 800 and 1000 milliosmol per kilogram and exceeded pi in the flexor by 300 to 450 milliosmol per kilogram. In fully closed pulvini the reverse was true, with pi in the flexor between 800 and 1000 milliosmol per kilogram, exceeding pi in the extensor by 300 to 450 milliosmol per kilogram. To obtain approximate values of Psi of pulvinar tissues, shallow cuts in extensor and flexor sides of oil-covered pulvini were filled with droplets of polyethylene glycol solutions of known Psi. Droplets maintaining constant size were assumed to have the same Psi as the tissue. Extensor and flexor halves of open pulvini had very different Psi (extensor, about -1.4 MPa; flexor, about -0.3 MPa), but both sides of closed pulvini had similar Psi (about -0.3 MPa). Measurements of Psi and pi and calculations of P indicate: (a) In open pulvini, P is about the same in extensor and flexor. The large Psi gradient is caused by a large osmotic gradient. (b) In closed pulvini, P is approximately 50% higher in the flexor than in the extensor. This difference in P compensates for differences in pi such that the Psi gradient is small. (c) Pulvini close as P increases in the flexor and reopen as flexor P decreases; extensor P values are similar in open and closed pulvini.

Journal Article↗
PubMed1990

Anatomical Changes and Immunolocalization of Cellulase during Abscission as Observed on Nitrocellulose Tissue Prints.

A fundamental event in abscission is the breakdown of cell wall material in a discrete zone of cells known as the separation layer. Three dimensional images produced by viewing tissue prints of abscission zones on nitrocellulose (NC) membranes with incident illumination showed changes in the tissue integrity taking place in the separation layer as the process of abscission proceeded. The cell softening which occurs due to the dissolution of the cell wall appeared in the tissue prints as a diffuse line at the anatomical transition between the pulvinus and petiole and was easily observed on NC tissue prints of either longitudinal or serial cross-sections through abscission zones. In bean leaf abscission the dissolution of cell walls has been correlated with the appearance of a form of cellulase with an isoelectric point of pH 9.5. Antibodies specific for this enzyme were used to study the localization of 9.5 cellulase in the distal abscission zone of Phaseolus vulgaris L., cv Red Kidney after tissue printing on NC. It was found that 9.5 cellulase was localized in the separation layer but also occurred in the vascular tissue of the adjacent pulvinus. No antibody binding was observed in nonabscising tissue or preimmune controls. These results confirm previous biochemical studies and demonstrate that immunostaining of nitrocellulose tissue prints is a fast and reliable method to localize proteins or enzymes in plant tissue.

Journal Article↗