An active role of the amyloplasts and nuclei of root statocytes in graviperception.

Three main phases are discerned in the gravitropic reaction: perception of a gravitational stimulus, its transduction, and fixation of the reaction resulting in bending of an organ. According to the starch-statolith hypothesis of Nemec and Haberlandt, amyloplasts in the structurally and functionally specialized graviperceptive cells (statocytes) sediment in the direction of a gravitational vector in the distal part of a cell while a nucleus is in the proximal one. If amyloplasts appear to act as gravity sensors, the receptors, which interact with sedimented amyloplasts, and next signaling are still unclear. An analysis of the structural-functional organization of cells in different root cap layers of such higher plants as pea, Arabidopsis thaliana, and Brassica rapa grown under 1 g, on the clinostats, and in microgravity, allows us to support the hypothesis that amyloplasts function as statoliths in statocytes, but they may not be only the passive statolithic mass. We propose that amyloplasts fulfill a more complex function by interacting with a receptor, which is a nucleus, in transduction of some signal to it. Gravity-induced statolith movement in certain order leads to a new functional connection between gravity susceptors--amyloplasts and a receptor--a nucleus receiving some signal presumedly of a mechanical or biochemical nature from the amyloplasts. During gravitropism, sugar signaling could induce expression of genes encoding auxin transport proteins in a nucleus giving the nucleus an intermediate role in signal trunsduction following perception.     

Transduction of the gravity stimulus in the root statocyte.

The amyloplasts of root statocytes are considered to be the perceptors of gravity. However, their displacement and the starch they contain are not required for gravisensing. The mechanism of the transduction of gravistimulus remains therefore controversial. It is well known that the amplitude of the stimulus is dependent upon the intensity of the acceleration and the inclination of the root with respect to gravity. This strongly supports the hypothesis that the stimulus results in a mechanical effect (pressure or tension) on a cellular structure. Three cellular components are proposed as possible candidates for the role of transducer: the actin filaments, the endoplasmic reticulum and the plasma membrane with its ion channels. Recent results obtained in the frame of the IML 1 Mission of Spacelab show that the endoplasmic reticulum should rather be responsible for the termination of the stimulus. The contacts of amyloplasts with the distal ER could therefore be involved in the regulation of root growth.     

Cell biology of plant gravity sensing.

The debate about whether gravity sensing relies upon statoliths (amyloplasts that sediment) has intensified with recent findings of gravitropism in starchless mutants and of claims of hydrostatic gravity sensing. Starch and significant plastid sedimentation are not necessary for reduced sensing in mutant roots, but plastids might function here if there were a specialized receptor for plastid mass e.g. in the ER. Alternatively, components in addition to amyloplasts might provide mass for sensing. The nucleus is dense and its position is regulated, but no direct data exist for its role in sensing. If the weight of the protoplast functioned in sensing, why would there be specific cytological specializations favoring sedimentation rather than cell mass? Gravity has multiple effects on plants in addition to gravitropism. There may be more than one mechanism of gravity sensing.     

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.     

Growth of pea epicotyl in low magnetic field implication for space research

A magnetic field is an inescapable environmental factor for plants on the earth. However, its impact on plant growth is not well understood. In order to survey how magnetic fields affect plant, Alaska pea seedlings were incubated under low magnetic field (LMF) and also in the normal geo-magnetic environment. Two-day-old etiolated seedlings were incubated in a magnetic shield box and in a control box. Sedimentation of amyloplasts was examined in the epicotyls of seedlings grown under these two conditions. The elongation of epicotyls was promoted by LMF. Elongation was most prominent in the middle part of the epicotyls. Cell elongation and increased osmotic pressure of cell sap were found in the epidermal cells exposed to LMF. When the gravitational environment was 1G, the epicotyls incubated under both LMF and normal geomagnetic field grew straight upward and amyloplasts sedimented similarly. However, under simulated microgravity (clinostat), epicotyl and cell elongation was promoted. Furthermore, the epicotyls bent and amyloplasts were dispersed in the cells in simulated microgravity. The dispersion of amyloplasts may relate to the posture control in epicotyl growth under simulated microgravity generated by 3D clinorotation, since it was not observed under LMF in 1G. Since enhanced elongation of cells was commonly seen both at LMF and in simulated microgravity, all elongation on the 3D-clinostat could result from pseudo-low magnetic field, as a by-product of clinorotation. (i.e., clinostat results could be based on randomization of magnetic field together with randomization of gravity vector.) Our results point to the possible use of space for studies in magnetic biology. With space experiments, the effects of dominant environmental factors, such as gravity on plants, could be neutralized or controlled for to reveal magnetic effects more clearly.     

The effects of HGMFs on the plant gravisensing system.

High Gradient Magnetic Fields (HGMFs) offer new opportunities for studying the gravitropic system of plants. However, it is necessary to analyze the influence that HGMF can have on cellular processes and structures that may not be related to amyloplasts displacement. This paper considers possible HGMF effects on plants, which may accompany HGMF stimulation of amyloplasts and contribute to the mechanisms of the HGMF-induced curvature.     

Perception of gravity by plants.

Physical principles can be used to predict some features about the gravity perception system in plants. The nature of the system has made it rather elusive, so this approach represents an additional source of information to help find it. For a gravitational stimulus to be detected, two masses must move relative to each other in a manner which causes a significant amount of work to be done on a receptor. Relative to cellular dimensions, the masses must be large, be dense and move noticeable distances. The main sources of noise are thermal motion and flexing of the plant tissue. Some new models for the function of amyloplasts as statoliths are presented.     

Gravitropism of axial organs in multicellular plants.

Gravitropism of plant organs such as roots, stems and coleoptiles can be separated into four distinct phases: 1. perception (gravity sensing), 2. transduction of a signal into the target region and 3. the response (differential growth). This last reaction is followed by a straightening of the curved organ (4.). The perception of the gravitropic stimulus upon horizontal positioning of the organ (1.) occurs via amyloplasts that sediment within the statocytes. This conclusion is supported by our finding that submerged rice coleoptiles that lack sedimentable amyloplasts show no graviresponse. The mode of signal transduction (2.) from the statocytes to the peripheral cell layers is still unknown. Differential growth (3.) consists of a cessation of cell expansion on the upper side and an enhancement of elongation on the lower side of the organ. Based on the facts that the sturdy outer epidermal wall (OEW) constitutes the growth-controlling structure of the coleoptile and that growth-related osmiophilic particles accumulate on the upper OEW, it is concluded that the differential incorporation of wall material (presumably glycoproteins) is causally involved. During gravitropic bending, electron-dense particles ('wall-loosening capacity') accumulate on the growth-inhibited upper OEW. It is proposed that the autotropic straightening response, which is in part due to an acceleration of cell elongation on the curved upper side, may be attributable to an incorporation of the accumulated particles ('release of wall-loosening capacity'). This novel mechanism of autotropic re-bending and its implications for the Cholodny-Went hypothesis are discussed.     

Sedimentable amyloplasts in starch sheath cells of woody stems of Japanese cherry.

We examined whether sedimentable amyloplasts act as statolith in the perception of gravity in woody stems using the elongated internodes of Japanese cherry (Prunus jamasakura Sieb. ex Koidz.). In the internode of the seedlings grown on earth, amyloplasts were found sedimented at the distal end of each cell of the endodermal starch sheath tissue. In the internode grown on three-dimensional (3-D) clinostat, amyloplasts were dispersed throughout the cell matrix in the endodermal starch sheath tissue. After changing the positions of the internode from vertical to horizontal, re-sedimentation of amyloplasts toward the direction of gravity was completed in 1h, whereas the bending of the internode was observed after 12 days. We propose that sedimentable amyloplasts in the endodermal starch sheath cells may play a role in gravity perception leading to secondary xylem formation in the secondary thickening growth and eccentric growth in gravi-bending of tree stems.     

新的像机观测可靠度模型

针对像机观测方向和观测区域受限的特点,结合人眼感知概率模型,建立了一种基于观测可靠度的像机观测模型.依据Johnson准则,综合考虑了像机距目标的观测距离、观测角度和目标大小等因素,不仅可以确定目标的存在与否,还可以全面描述像机对目标的观测性能,更加符合像机对目标感知的实际情况.在此基础上定义了多个像机对目标的观测可靠度,及像机网络对目标的捕获概率,提出了基于像机观测可靠度模型的多像机网络覆盖优化部署算法.仿真结果表明,利用新的像机观测可靠度模型对网络进行优化,可以在提高多像机网络对观测区域的覆盖程度的同时,优化网络对场景的感知能力,提高对目标的捕获概率.      

Laser ablation of root cap cells: implications for models of graviperception.

The initial event of gravity perception by plants is generally thought to occur through sedimentation of amyloplasts in specialized sensory cells. In the root, these cells are the columella which are located toward the center of the root cap. To define more precisely the contribution of columella cells to root gravitropism, we used laser ablation to remove single columella cells or groups of these cells and observed the effect of their removal on gravity sensing and response. Complete removal of the cap or all the columella cells (leaving peripheral cap cells intact) abolishes the gravity response of the root. Removal of stories of columella revealed differences between regions of the columella with respect to gravity sensing (presentation time) versus graviresponse (final tropic growth response of the root). This fine mapping revealed that ablating the central columella located in story 2 had the greatest effect on presentation time whereas ablating columella cells in story 3 had a smaller or no effect. However, when removed by ablation the columella cells in story 3 did inhibit gravitropic bending, suggesting an effect on translocation of the gravitropic signal from the cap rather than initial gravity perception. Mapping the in vivo statolith sedimentation rates in these cells revealed that the amyloplasts of the central columella cells sedimented more rapidly than those on the flanks do. These results show that cells with the most freely mobile amyloplasts generate the largest gravisensing signal consistent with the starch statolith hypothesis of gravity sensing in roots.     

航空发动机整机有限元模型转子动力学分析

现代航空发动机在工作中不断变化的机械激振,气动激振频率越来越复杂,这使得对航空发动机振动分析必须考虑各结构间的动力影响.因此,利用能够考虑陀螺力矩影响,基于NASTRAN中实体单元编制的转子动力特性计算程序,对发动机整机进行了动力特性计算.首先对转子支承结构传递函数(动刚度)进行计算,并进一步研究其对转子动力特性的影响;分析比较基于不同单元模型计算时,盘轴耦合振动及盘轴连接处的角刚度对转子动力特性的影响,证明了基于实体单元的整机模型能够准确考虑各种振动模态.最后,在分析中发现了高阶转子弯曲振动模态与机匣振动耦合现象及其变化规律,在计算分析的基础上研究了在考虑机匣振动耦合时转子系统临界转速的确定方法.      

从空间飞行器入射进电离层的调制电子束产生的高频波辐射

在主动束-等离子体试验中,调制电子束从空间飞行器入射进电离层等离子体将会产生电磁波辐射,在不同试验条件下电磁波辐射机理也不一样,由电子束纵向约束性产生电磁波辐射是其中之一.对半无界稀薄调制电子束从空间飞行器入射进电离层等离子体时所产生的波现象进行了理论分析和数值计算.结果表明,当调制电子束沿磁力线入射时,会在电离层等离子体中产生高频电磁波辐射,该辐射主要集中在垂直于入射电子束运动方向的平面内.      

Mechanical analysis of statolith action in roots and rhizoids.

Published observations on the response times following gravistimulation (horizontal positioning) of Chara rhizoids and developing roots of vascular plants with normal and "starchless" amyloplasts were reviewed and compared. Statolith motion was found to be consistent with gravitational sedimentation opposed by elastic deformation of an intracellular material. The time required for a statolith to sediment to equilibrium was calculated on the basis of its buoyant density and compared with observed sedimentation times. In the examples chosen, the response time following gravistimulation (from horizontal positioning to the return of downward growth) could be related to the statolith sedimentation time. Such a relationship implies that the transduction step is rapid in comparison with the perception step following gravistimulation of rhizoids and developing roots.     

Magnetospheric Boundary Layer Structure and Dynamics as Seen From Cluster and Double Star Measurements

In this review, we discuss the structure and dynamics of the magnetospheric Low-Latitude Boundary Layer (LLBL) based on recent results from multi-satellite missions Cluster and Double Star. This boundary layer, adjacent to the magnetopause on the magnetospheric side, usually consists of a mixture of plasma of magnetospheric and magnetosheath origins, and plays an important role in the transfer of mass and energy from the solar wind into the magnetosphere and subsequent magnetospheric dynamics. During southward Interplanetary Magnetic Field (IMF) conditions, this boundary layer is generally considered to be formed as a result of the reconnection process between the IMF and magnetospheric magnetic field lines at the dayside magnetopause, and the structure and plasma properties inside the LLBL can be understood in terms of the time history since the reconnection process. During northward IMF conditions, the LLBL is usually thicker, and has more complex structure and topology. Recent observations confirm that the LLBL observed at the dayside can be formed by single lobe reconnection, dual lobe reconnection, or by sequential dual lobe reconnection, as well as partially by localized cross-field diffusion. The LLBL magnetic topology and plasma signatures inside the different sub-layers formed by these processes are discussed in this review. The role of the Kelvin-Helmholtz instability in the formation of the LLBL at the flank magnetopause is also discussed. Overall, we conclude that the LLBL observed at the flanks can be formed by the combination of processes, (dual) lobe reconnection and plasma mixing due to non-linear Kelvin-Helmholtz waves.      

The Neupert effect and new RHESSI measures of thetotal energy in electrons accelerated in solar flares

It is believed that a large fraction of the total energy released in a solar flare goes initially into acceleratedelectrons. These electrons generate the observed hard X-ray bremsstrahlung as they lose most of their energy by coulomb collisions in the lower corona and chromosphere. Results from the Solar Maximum Mission showed that there may be even more energy in accelerated electrons with energies above 25 keV than in the soft X-ray emitting thermal plasma. If this is the case, it is difficult to understand why the Neupert Effect — the empirical result that for many flares the time integral of the hard X-ray emission closely matches the temporal variation of the soft X-ray emission — is not more clearly observed in many flares. From recent studies, it appears that the fraction of the released energy going into accelerated electrons is lower, on average, for smaller flares than for larger flares. Also, from relative timing differences, about 25% of all flares are inconsistent with the Neupert Effect. The Reuven Ramaty High Energy Solar Spectroscopic Imager (RHESSI) is uniquely capable of investigating the Neupert Effec since it covers soft X-rays down to 3 keV (when both attenuators are out of the field of view) and hard X-rays with keV energy resolution, arcsecond-class angular resolution, and sub-second time resolution. When combined with the anticipated observations from the Soft X-ray Imager on the next GOES satellite, these observations will provide us with the ability to track the Neupert Effect in space and time and learn more about the relation between plasma heating and particle acceleration. The early results from RHESSI show that the electron spectrum extends down to as low as 10 keV in many flares, thus increasing the total energy estimates of the accelerated electrons by an order of magnitude or more compared with the SMM values. This combined with the possible effects of filling factors smaller than unity for the soft X-ray plasma suggest that there is significantly more energy in nonthermal electrons than in the soft X-ray emitting plasma in many flares.     

太阳软X射线背景水平与X级的增强发射

本文根据1986年1月-1992年12月期间,130个1-8?的X级别的X射线事件与相应背景发射水平所作的分析,发现90%以上的X级X射线事件,都发生在X射线背景发射≥C1(10-6W／m2)水平上,这一统计规律,可从耀斑加速的非热电子对色球等离于体的加热得到合理的解释.      

Polycyclic aromatic hydrocarbons: primitive pigment systems in the prebiotic environment.

Polycyclic aromatic hydrocarbons (PAH) in the form of polymerized derivatives represent over 90% of the organic material of carbonaceous chondrites. It now appears likely that there was substantial survival of the organic content of meteoritic and cometary infall during late accretion, so that PAH would presumably be major components of the organic inventory present on the prebiotic Earth. An important question relative to chemical evolution and energy transduction is the nature of pigments which could be available to make light energy available to the earliest cellular forms of life. PAH and their derivatives all absorb light in the near UV and blue wavelengths, and are candidates for primitive pigments. We have explored this possibility in a model system consisting of mixtures of pyrene, fluoranthene and pyrene derivatives with hexadecane, dispersed in dilute salt solutions. Upon illumination, photochemical oxidation of the hexadecane occurs, with long-chain amphiphiles such as 2-hexadecanone and 2-hexadecanol as products. Because the reaction proceeds under strictly anaerobic conditions, the source of oxygen is apparently water. We also observed acid pH shifts during illumination. Photochemical production of hydrogen ion is significant, in that chemiosmotic proton gradients across membranes are used by all contemporary cells as a source of energy for ATP synthesis and nutrient transport. To test whether the protons could be used to transduce light energy into a useful form, PAH derivatives were included in lipid bilayer membranes (liposomes). Upon illumination, protons (or acidic products) were produced and accumulated inside the vesicles, so that substantial pH gradients were established across the membranes, acid inside. We conclude that PAH dissolved in aliphatic hydrocarbons absorb light energy and use it to oxidize the hydrocarbon to long-chain amphiphilic molecules. The oxidation is accompanied by release of protons. If PAH derivatives are included in the bilayer membrane of lipid vesicles, protons accumulate within the membrane-bounded volumes to form proton gradients. This system provides a useful model of a primitive photochemical reaction in which light energy is transduced into potentially useable forms.     

The relationship between profiles of plagiogravitropism and morphometry of columella cells during the development of lateral roots of Vigna angularis

There has been no convincing explanation on a mechanism inducing plagiogravitropism of lateral roots. The present work deals with gravitropic features of Vignaangularis lateral roots during the course of their growth and morphometric analysis of root caps, columella cells and amyloplasts. Regardless of the magnitude of deviation of the primary root axis from the gravity vector, the newly emerging lateral roots tended to keep a constant angle to the gravity vector. They modified gravireaction several times during the course of their development: a first horizontal-growth stage when they grow in the cortex of primary roots (stage I), a sloping-down growth stage from their emergence to a length of about 1 mm (stage II), a second horizontal-growth stage from a length of about 1 mm to that of over 4 mm (stage III) and a curving-down stage thereafter (stage IV). The columella cells with amyloplasts large enough to sediment were not fully differentiated in the stage I but the turning point from the stage I to II was associated with the development of amyloplasts which were able to sediment toward the distal part of the cell. Amyloplasts were significantly small in the lateral roots over 10 mm long compared with those in ones 0–10 mm long, suggesting that they rapidly develop immediately after the lateral roots emerge from primary roots and then gradually decrease their size when the lateral roots grow over 10 mm long. This dimensional decrease of amyloplasts may be partially involved in weak gravireaction in the stage III. Evidence was not presented indicating that a switchover from the stage III to IV was connected with the dimension of root caps, the number of columella cells and the development of amyloplasts. Some factors at the molecular level rather than at the cellular and tissue levels are probably dominant to induce the stage IV.     

Probing heliospheric diffusion coefficients with solar energetic particles

The dynamics of solar particle events provide a direct link to the understanding of properties of wave–particle interactions, and to the nature of the solar wind fluctuations. Depending on their energy, the often simultaneously observed electrons, protons and ions interact with different wavenumber ranges of the fluctuations, and are sensitive to various aspects of the dynamical nature of the solar wind turbulence. In general, the evolution of particle events is also sensitive to the spatial variation of the transport parameters between the Sun and a few AU. Together with in situ plasma and magnetic field observations this information can be used to extrapolate the properties of transport parameters into the more distant Heliosphere. Recent developments in the theory of parallel transport of energetic particles, and examples for the modelling of solar particle events and the derivation of transport parameters are considered. A dynamical quasi-linear theory is presented which gives special emphasis to the geometry and dynamic nature of the fluctuations, and which is able to provide particle mean free paths solely from observed plasma parameters, in good agreement with those derived by the modelling. Possibilities to apply the above results to the study of other energetic particle processes in the Heliosphere are discussed.