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PubMed1975

Abscission of Phaseolus and Impatiens Explants: Effects of Ionizing Radiation upon Endogenous Growth Regulators and de Novo Enzyme Synthesis.

Stem-petiole explants from the lower pulvinus of the primary leaves of Phaseolus vulgaris L. cv. Red Kidney and from Impatiens sultani Hook cv. Scarlet Baby were exposed to varying dosages of gamma-radiation. With bean, irradiation of 175 to 525 kiloroentgens (kR) significantly accelerated the onset of abscission with a maximum response at 175 to 280 kR. Higher dosages (beginning at 600-700 kR) usually prevented abscission. With Impatiens, 18 to 35 kR significantly accelerated both the onset of abscission and possibly the initial abscission rate; 350 kR cut the time to 100% abscission in half and substantially accelerated the initial abscission rate. Inhibition of abscission in Impatiens was not possible with the available dose rate (35 kR/hour).The accelerating effect of ionizing radiation was apparently due to primarily a reduction in auxin activity, thus accelerating the shift from the relatively ethylene-insensitive stage I to the ethylene-sensitive stage II. In vitro irradiation of exogenous auxin in agar substantially reduced the abscission-delaying effects of the auxin. The low dosage accelerating effects of the ionizing radiation were negated by exogenous auxin application, either prior to or following irradiation. With prior auxin application, a higher concentration of auxin was required to achieve the same delaying effects as with application following irradiation. Apparently due to this reduction in auxin activity, irradiation hastened sensitivity to exogenous ethylene.The accelerating effects of ionizing radiation may be secondarily due to increased levels of endogenous ethylene. With bean tissues, exposures of 262 through 1500 kR of gamma radiation increased the measurable levels of endogenous ethylene released into the ambient air within 2 or 3 days after irradiation.The results of actinomycin D treatment suggested that required de novo RNA synthesis occurred about 4 hours earlier in those Impatiens explants that had received 105 kR gamma radiation than in the nonirradiated controls.An increased mobilization of Ca from the abscission zone may be a contributing factor in the radiation-induced acceleration of abscission. For both bean and Impatiens, exogenous Ca inhibited abscission of irradiated and nonirradiated explants. In addition, a more concentrated Ca solution was required to inhibit abscission of irradiated bean explants, as compared to the nonirradiated controls.

Journal Article↗
PubMed1983

H fluxes in excised samanea motor tissue : I. Promotion by light.

Previous investigators revealed that white light-promoted leaflet opening in Samanea saman (Jacq) Merrill depends upon K(+) uptake by extensor cells and efflux from flexor cells of the pulvinus, while dark-promoted closure depends upon K(+) fluxes in the opposite directions. We now monitored H(+) fluxes during pulvinar movement to test a model proposing coupled H(+)/K(+) fluxes. H(+) fluxes were monitored by measuring changes in the pH of a weakly buffered solution (initial pH = 5.5) bathing excised strips of extensor or flexor tissue. White light at hour 3 of the usual dark period promoted pulvinar opening, H(+) efflux from extensor cells and uptake by flexor cells, while darkness at hours 2 to 4 of the usual light period promoted pulvinar closure, H(+) uptake by extensor cells and efflux from flexor cells. The following conditions altered H(+) fluxes during dark-promoted closure. (a) Light reversed the directions of the fluxes in both extensor and flexor cells. (b) Anoxia increased the rate of H(+) uptake by extensor cells and promoted H(+) uptake (rather than efflux) by flexor cells, consistent with an outwardly directed H(+) pump. KCN showed similar effects initially, but they were transient. (c) Increase in external pH from 5.5 to 6.7 promoted H(+) efflux (rather than uptake) by extensor cells and increased the rate of H(+) efflux from flexor cells, presumably by decreasing the rate of inward diffusion. (d) Change in external K(+) did not alter H(+) fluxes by extensor cells, but removal of external K(+) decreased the rate of H(+) efflux from flexor cells by 70%. These observations support a model for coupled H(+)/K(+) fluxes in pulvinar cells during light-and dark-promoted leaflet movements.

Journal Article↗
PubMed1984

Deferral of senescence and abscission by chemical inhibition of ethylene synthesis and action in bean explants.

Three compounds known to inhibit ethylene synthesis and/or action were compared for their ability to delay senescence and abscission of bean explants (Phaseolus vulgaris L. cv Contender). Aminoethoxyvinyl-glycine (AVG), AgNO(3), and sodium benzoate were infiltrated into the petiole explants. Their effect on abscission was monitored by measuring the force required to break the abscission zone, and their effect on senescence was followed by measuring chlorophyll and soluble protein in the distal (pulvinus) sections. AVG at concentrations between 1 and 100 micromolar inhibited ethylene synthesis by about 80 to 90% compared to the control during sampling periods of 24 and 48 hours after treatment. This compound also delayed the development of abscission and senescence. Treatment with AgNO(3) at concentrations between 1 and 100 micromolar progressively reduced ethylene production, but to a lesser extent than AVG. The effects of AgNO(3) on senescence and abscission were quite similar to those of AVG. Sodium benzoate at 50 micromolar to 5 millimolar did not inhibit ethylene synthesis during the first 24 hours, but appreciably inhibited ethylene synthesis 48 hours after treatment. It also delayed the development of abscission and senescence. The effects of AVG, Ag(+), and sodium benzoate suggest that ethylene could play a major role in both the senescence induction phase and the separation phase in bean explants.

Journal Article↗
PubMed1986

Diurnal Phototropism in Solar Tracking Leaves of Lavatera cretica.

On a clear day, leaf laminas of Lavatera cretica tracked the solar position throughout the day. The laminar azimuth did not diverge from the solar azimuth by more than 12 degrees from sunrise to sunset. Tracking of the solar elevation started 1 to 2 hours after sunrise and ceased 1 to 2 hours before sunset. On an overcast day, the laminas reoriented horizontally. After sunset, following a clear day, the laminas performed a nocturnal reorientation, with three well defined phases. During the initial phase the laminas relaxed their strained sunset-facing orientation to one perpendicular to their petioles. This equilibrium configuration was maintained throughout the following phase, which was apparently concerned with time-measuring. During the final phase, the laminas reoriented, before sunrise, to a position facing the direction of the anticipated sunrise. This directional information is phototropic and was retained for 3 to 4 diurnal cycles, probably in the pulvinus itself, which is the site of the response. Laminas of plants transferred from sunlight either to darkness, or to a simulated natural photoperiod under overhead illumination, were facing the originally anticipated direction of sunrise at the time of each of the three to four subsequent sunrises (after which they reverted to the dark orientation in darkness, or to the horizontal one with overhead illumination). Cotyledonary laminas required directional information for the nocturnal reorientation during 3 or 4 cycles of simulated sunrise to sunset transitions.

Journal Article↗
PubMed1987

Gibberellins and gravitropism in maize shoots: endogenous gibberellin-like substances and movement and metabolism of [3H]Gibberellin A20.

[3H]Gibberellin A20 (GA20) of high specific radioactivity (49.9 gigabecquerel per millimole) was applied equilaterally in a ring of microdrops to the internodal pulvinus of shoots of 3-week-old gravistimulated and vertical normal maize (Zea mays L.), and to a pleiogravitropic (prostrate) maize mutant, lazy (la). All plants converted the [3H]GA20 to [3H]GA1- and [3H]GA29-like metabolites as well as to several metabolites with the partitioning and chromatographic behavior of glucosyl conjugates of [3H]GA1, [3H]GA29, and [3H]GA8. The tentative identification of these putative [3H]GA glucosyl conjugates was further supported by the release of the free [3H]GA moiety after cleavage with cellulase. Within 12 hours of the [3H]GA20 feed, there was a significantly higher proportion of total radioactivity in lower than in upper halves of internode and leaf sheath pulvini in gravistimulated normal maize. Further, there was a significantly higher proportion of putative free GA metabolites of [3H]GA20, especially [3H]GA1, in the lower halves of normal maize relative to upper halves. The differential localization of the metabolites between upper and lower halves was not apparent in the pleiogravitropic mutant, la. Endogenous GA-like substances were also examined in gravistimulated maize shoots. Forty-eight hours after gravistimulation of 3-week-old maize seedlings, endogenous free GA-like substances in upper and lower leaf sheath and internode pulvini halves were extracted, chromatographed, and bioassayed using the "Tanginbozu" dwarf rice microdrop assay. Lower halves contained consistently higher total levels of GA-like activity. The qualitative elution profile of GA-like substances differed consistently, upper halves containing principally a GA20-like substance and lower halves containing principally a GA20-like substance and lower halves containing mainly GA1-like and GA19-like substances. Gibberellins A1 (10 nanograms per gram) and A20 (5 nanograms per gram) were identified from these lower leaf sheath pulvini by capillary gas chromatography-selected ion monitoring. Results from all of these experiments are consistent with a role for GAs in the differential shoot growth that follows gravitropism, although the results do not eliminate the possibility that the redistribution of GAs results from the gravitropic response.

Gibberellins↗
PubMed1988

Potassium Channels in Motor Cells of Samanea saman: A Patch-Clamp Study.

Leaflet movements in Samanea saman are driven by the shrinking and swelling of cells in opposing (extensor and flexor) regions of the motor organ (pulvinus). Changes in cell volume, in turn, depend upon large changes in motor cell content of K(+), Cl(-) and other ions. We performed patch-clamp experiments on extensor and flexor protoplasts, to determine whether their plasma membranes contain channels capable of carrying the large K(+) currents that flow during leaflet movement. Recordings in the "whole-cell" mode reveal depolarization-activated K(+) currents in extensor and flexor cells that increase slowly (t((1/2)) = ca. 2 seconds) and remain active for minutes. Recordings from excised patches reveal a single channel conductance of ca. 20 picosiemens in both cell types. The magnitude of the K(+) currents is adequate to account quantitatively for K(+) loss, previously measured in vivo during cell shrinkage. The K(+) channel blockers tetraethylammonium (5 millimolar) or quinine (1 millimolar) blocked channel opening and decreased light- and dark-promoted movements of excised leaflets. These results provide evidence for the role of potassium channels in leaflet movement.

Journal Article↗
PubMed1988

Changes in Two Forms of Membrane-Associated Cellulase during Ethylene-Induced Abscission.

Only one form of membrane-associated cellulase was found previously in the lower petiolar pulvinus of Phaseolus vulgaris (cv Red Kidney). The cellulase has an isoelectric point (pI) of 4.5 (DE Koehler, LN Lewis 1979 Plant Physiol 63: 677-679). This enzyme was detected in abscission zones collected before the onset of abscission (control tissue), and was thought to represent a pre-secretory form of another cellulase, the abscission cellulase, which has a basic pI and is secreted during abscission. We now show that this acidic, membrane-associated cellulase is a glycoprotein, tightly bound to the membrane, with maximum activity at pH 5.1, and that it is not immunologically related to the abscission cellulase. Furthermore, when bean explants are induced to abscise with ethylene, the activity of the acidic cellulase declines rapidly to 50% of control levels in the first day. When abscission is fully developed, the membranes contain a basic form of cellulase with a pI of 8.0 to 9.0 and only trace levels of the acidic cellulase. The basic form is not a high mannose glycoprotein; it has maximum activity in a broad pH range (4.0-8.0) and is antigenically related to the abscission cellulase, which is induced during abscission and transported to the cell wall. Antibody raised against the abscission cellulase recognized two proteins in a crude membrane fraction from abscising tissue. One of those proteins comigrated with the abscission cellulase, and the other was 1 to 2 kilodaltons larger. Thus, during abscission, the acidic membrane-associated cellulase rapidly declines before the appearance of the abscission cellulase. We conclude that there is no conversion from the acidic cellulase to the basic cellulase and suggest that the acidic and basic cellulase isoenzymes are proteins derived from two different genes.

Journal Article↗
PubMed1992

Effects of Light on the Membrane Potential of Protoplasts from Samanea saman Pulvini : Involvement of K Channels and the H -ATPase.

Rhythmic light-sensitive movements of the leaflets of Samanea saman depend upon ion fluxes across the plasma membrane of extensor and flexor cells in opposing regions of the leaf-movement organ (pulvinus). We have isolated protoplasts from the extensor and flexor regions of S. saman pulvini and have examined the effects of brief 30-second exposures to white, blue, or red light on the relative membrane potential using the fluorescent dye, 3,3'-dipropylthiadicarbocyanine iodide. White and blue light induced transient membrane hyperpolarization of both extensor and flexor protoplasts; red light had no effect. Following white or blue light-induced hyperpolarization, the addition of 200 millimolar K(+) resulted in a rapid depolarization of extensor, but not of flexor protoplasts. In contrast, addition of K(+) following red light or in darkness resulted in a rapid depolarization of flexor, but not of extensor protoplasts. In both flexor and extensor protoplasts, depolarization was completely inhibited by tetraethylammonium, implicating channel-mediated movement of K(+) ions. These results suggest that K(+) channels are closed in extensor plasma membranes and open in flexor plasma membranes in darkness and that white and blue light, but not red light, close the channels in flexor plasma membranes and open them in extensor plasma membranes. Vanadate treatment inhibited hyperpolarization in response to blue or white light, but did not affect K(+) -induced depolarization. This suggests that white or blue light-induced hyperpolarization results from activation of the H(+) -ATPase, but this hyperpolarization is not the sole factor controlling the opening of K(+) channels.

Journal Article↗
PubMed1990

Occurrence and Localization of 9.5 Cellulase in Abscising and Nonabscising Tissues.

Nitrocellulose tissue prints immunoblotted with 9.5 cellulase antibody were used to demonstrate areas of cellulase localization within Phaseolus vulgaris explants on exposure to ethylene. The 9.5 cellulase was induced in the distal and proximal abscission zone and in the stem. In both abscission zones, the 9.5 cellulase was found in the cortical cells of the separation layer, which develops as a narrow band of cells at the place where fracture occurs. The enzyme was also found associated with the vascular traces of the tissues adjacent to the separation layer extending through the first few millimeters at each side of the separation layer. The two abscission zones differed in the way that cellulase distributed through the separation layer as abscission proceeded. In the distal zone, cellulase appeared first in the cells of the separation layer adjacent to vascular traces and extended toward the periphery. In the proximal zone, 9.5 cellulase accumulated first in the cortical cells that lie in the adaxial side and then extended to the abaxial side. In response to ethylene, 9.5 cellulase was also induced in the vascular traces of the stem and the pulvinus without developing a separation layer. The role of 9.5 cellulase in the vascular traces is unknown. Sodium dodecyl sulfate-polyacrylamide gel electrophoresis followed by immunoblotting with 9.5 cellulase antibody identified the same 51-kilodalton protein in both abscising and nonabscising tissues. Therefore, the determinant characteristic of the abscission process is the induction of 9.5 cellulase by cortical cells in the separation layer, and this implies that these cells have a unique mechanism for initiating 9.5 cellulase synthesis.

Journal Article↗
PubMed1988

An experimental analysis and comparison of three rhythms of movements in bean (Phaseolus vulgaris L.).

Three types of rhythmic movements of Phaseolus vulgaris L. (pole beans) were examined collectively and their characteristics compared. Although the ultradian rhythms of shoot circumnutation and leaf movement, as well as the circadian rhythm of leaf movement, occurred simultaneously, each rhythm could be expressed independently of the other two. Shoot circumnutation and ultradian leaf movements displayed the same period (80 min at 25 degrees C and Q10 congruent to 2), while the period of the circadian leaf movements was not temperature dependent (Q10 congruent to 1). Interaction into the plant between two ultradian rhythms (shoot circumnutation and ultradian leaf movement) with the same period and coexistence in the pulvinus of an ultradian with a circadian rhythm are discussed.

Activity Cycles↗
PubMed1968

Cell wall dissolution and enzyme secretion during leaf abscission.

Cell wall changes leading to the formation of the separation layer during abscission of unifoliate (Phaseolus vulgaris) leaves are reviewed. Based on evidence from explants and intact plants, dissolution of pectic substances between cells of adjacent tissue regions (petiole and pulvinus) is necessary and may be sufficient to form the separation layer. Initially, the abscission zone is not structurally weak. The decline in break strength accompanying formation of the separation layer correlates with the appearance of pectinase activity. Pectinase activity is not detectable in freshly harvested explants but increases to about 0.09 mug per abscission zone at the time of 50% separation. At the same time, water extractable pectin fractions increase with a corresponding decline in the pectin fraction extractable with dilute acid. Separation is aided by internal shear forces generated by differential growth and hydrostatic pressure or both.

Journal Article↗
PubMed1995

Calcium-sensitivity of the plasmalemmal delayed rectifier potassium current suggests that calcium influx in pulvinar protoplasts from Mimosa pudica L. can be revealed by hyperpolarization.

Isolated protoplasts from pulvinar motor cells of Mimosa pudica were studied using conventional whole-cell patch clamp techniques. With internal solutions weakly buffered for Ca2+ (0.2 mM EGTA), a rundown of the outward delayed rectifier K+ current was induced by hyperpolarizing the holding potential, and this effect was strongly promoted by high external Ca2+ concentrations. This rundown could be reversed by coming back to less hyperpolarized holding potentials or by lowering the external [Ca2+]. Such rundown was absent when pipette internal solutions strongly buffered (10 mM EGTA) for Ca2+ were used. Ionomycin induced rundown of the K+ current with internal solutions containing 0.2 mM but not 10 mM EGTA. The hyperpolarization-associated rundown was reversibly blocked by Gd3+ and La3+.

Bepridil↗
PubMed2001

Intracellular localization of phytochrome in Robinia pseudoacacia pulvini.

The intracellular localization of phytochrome in the pulvini of Robinia pseudoacacia L. was analyzed by immunogold electron microscopy after red (R; 15 min) and far-red (FR; 5 min) irradiation 2 h after the beginning of the photoperiod. Screening of the available antibodies by immunoblotting demonstrated that none of the oat (Avena sativa L.) anti-phytochrome A (phy A) monoclonal antibodies) (MAbs) detected Robinia phytochrome. A putative Robinia phy A was detected by immunoblotting using a MAb to mustard (Sinapis alba L.) phy A (CP 2/9). No cross-reactivity was observed in blots probed with a MAb against Cucumis sativus L. phy B (mAT1). Ultrathin sections of LR White resin-embedded pulvini were immunolabelled with CP 2/9 MAb. The labelling was restricted to cortical cells and there was no evidence of labelling either in the vascular system or in the epidermis. The pattern of labelling was the same in both extensor and flexor cells irrespective of whether phytochrome was in the far-red-absorbing (Pfr) state or had reverted to the red-absorbing (Pr) form. Isolated labels and clusters of labels were randomly distributed throughout the cytoplasm. Gold particles were also found in the interior of nuclei, chloroplasts and mitochondria.

Biological Transport↗
PubMed2006

Microautoradiographic localisation of [3H]sucrose and [3H]mannitol in Robinia pseudoacacia pulvinar tissues during phytochrome-mediated nyctinastic closure.

We have analysed the incorporation of [(3)H]sucrose and [(3)H]mannitol in pulvinar motor cells of Robinia pseudoacacia L. during phytochrome-mediated nyctinastic closure. Pairs of leaflets, excised 2 h after the beginning of the photoperiod, were fed with 50 mM [(3)H]sucrose or [(3)H]mannitol, irradiated with red (15 min) or far-red (5 min) light and placed in the dark for 2-3 h. Label uptake was measured in whole pulvini by liquid scintillation counting. The distribution of labelling in pulvinar sections was assessed by both light and electron microautoradiography. [(3)H]Sucrose uptake was twice that of [(3)H]mannitol incorporation in both red- and far-red-irradiated pulvini. In the autoradiographs, [(3)H]sucrose and [(3)H]mannitol labelling was localised in the area from the vascular bundle to the epidermis, mainly in vacuoles, cytoplasm, and cell walls. Extensor and flexor protoplasts displayed a different distribution of [(3)H]sucrose after red and far-red irradiation. Far-red light drastically reduced the [(3)H]sucrose incorporation in extensor protoplasts and caused a slight increase in internal flexor protoplasts. After red light treatment, no differences in [(3)H]sucrose labelling were found between extensor and flexor protoplasts. Our results indicate a phytochrome control of sucrose distribution in cortical motor cells and seem to rule out the possibility of sucrose acting as an osmoticum.

Autoradiography↗
PubMed1989

Plant movements caused by differential growth--unity or diversity of mechanisms?

A very wide range of plant organ movements have been described and yet it is not clear how each of them is related to the others. This uncertainty has had two undesirable consequences. Firstly, some workers have accepted the idea of a vague unity in the area and have subsequently been misled by information obtained in one system and applied without adequate justification to a study of a quite different system. Secondly, some researchers have evoked a possible diversity to explain why a particular mechanistic explanation may continue to be valid even when the model fails to explain events of a very similar nature in a slightly different system. We argue that this confusion has resulted from a classification of organ movements which has been based on functional rather than on mechanistic considerations. Mechanistic unity is to be expected on evolutionary grounds. This unity, however, may apply only to certain elements of the stimulus-response chain, at certain levels of organization. It follows from this that in seeking this unity, comparisons should be made between equivalent elements of the stimulus-response chain at the same level of organization in different systems. Only when this is done will theories built around the concept of unity provoke meaningful discussion.

Gravitropism↗
PubMed1995

Basis for changes in the auxin-sensitivity of Avena sativa (oat) leaf-sheath pulvini during the gravitropic response.

During the gravitropic response, auxin-sensitivity of the lower flanks of leaf-sheath pulvini of Avena sativa (oat) is at least 1000-fold higher than those of the upper flanks and non-gravistimulated pulvini. When the pulvini are treated with 1 mM Ca2+, a 10-fold increase in auxin-sensitivity of the pulvini is observed. Related to this difference in auxin-sensitivity, in vitro activation of the vanadate-sensitive H(-)-ATPase by IAA was observed. Results show that the activation of the H(+)-ATPase by IAA is probably mediated by soluble protein factors and that the H(+)-ATPase prepared from the lower flanks is activated by IAA with a 1000-fold higher auxin-sensitivity as compared with that from the upper flanks of the graviresponding pulvini. Ammonium sulfate fractionation experiments show that these soluble protein factors are in the 30 to 60% fraction. Auxin-binding assays reveal that lower flanks contain more high-affinity soluble auxin-binding sites (kD; on the order of 10(-9) M) and less low-affinity soluble auxin-binding sites (kD; on the order of 10(-6) M) than upper flanks. It is concluded that differential auxin-sensitivity of graviresponding oat-shoot pulvini is achieved by the modulation of affinities of auxin-binding sites in upper and lower flanks of the pulvini, that Ca2+ is involved in such modulation, and that one of the probable cellular functions of these auxin binding sites is the activation of the proton pump on the plasma membranes.

Avena↗
PubMed2003

Gravisensing: ionic responses, cytoskeleton and amyloplast behavior.

In Zea mays L., changes in orientation of stems are perceived by the pulvinal tissue, which responds to the stimulus by differential growth resulting in upward bending of the stem. Gravity is perceived in the bundle sheath cells, which contain amyloplasts that sediment to the new cell base when a change in the gravity vector occurs. The mechanism by which the mechanical signal is transduced into a physiological response is so far unknown for any gravity perceiving tissue. It is hypothesized that this involves interactions of amyloplasts with the plasma membrane and/or ER via cytoskeletal elements. To gain further insights into this process we monitored amyloplast movements in response to gravistimulation. In a pharmacological approach we investigated how the dynamics of plastid sedimentation are affected by actin and microtubule (MT) disrupting drugs. Dark grown caulonemal filaments of the moss Physcomitrella patens respond to gravity vector changes with a reorientation of tip growth away from the gravity vector. MT distributions in tip cells were monitored over time and MTs were seen to accumulate preferentially on the lower flank of the tip 30 min after a 90 degree turn. Using a self-referencing Ca2+ selective ion probe, we found that growing caulonemal filaments exhibit a Ca2+ influx at the apical dome, similar to that reported previously for other tip growing cells. However, in gravistimulated Physcomitrella filaments the region of Ca2+ influx is not confined to the apex, but extends about 60 micrometers along the upper side of the filament. Our results indicate an asymmetry in the Ca2+ flux pattern between the upper and side of the filament suggesting differential activation of Ca2+ permeable channels at the plasma membrane.

Actin Cytoskeleton↗
PubMed2000

Effects of staurosporine, okadaic acid and sodium fluoride on protein phosphorylation in graviresponding oat shoot pulvini.

This study focuses on the characterization of protein phosphorylation in the gravitropic response in oat shoot pulvini through the use of inhibitors of this process, namely staurosporine, okadaic acid and sodium fluoride. These three inhibitors reduce gravitropic curvature and cause changes in the phosphorylation of 38- and 50-kDa soluble proteins which show different levels of phosphorylation between lower and upper halves of gravistimulated pulvini. A kinetic analysis of phosphorylation shows that the 38- and 50-kDa soluble proteins exhibit different levels of phosphorylation between lower and upper halves of graviresponsive pulvini at 5 min after initiation of gravistimulation of stems. In addition, the phosphorylation of 63- and 70-kDa proteins from a total membrane preparation increases in lower halves of the pulvini following gravistimulation. These phosphoproteins are not found in the plasma membrane fraction. Taken together, at least four kinds of phosphoproteins are gravi-related. Of these, the 38- and 50-kDa soluble phosphoproteins may be involved in the regulation of early stages of the gravitropic response.

Avena↗