Anisotropy of wave velocities in plants: gravitropism

New measurements of W-wave velocities in plants suggest that the previously observed velocity of 96 cm/s in plants is not unique but that there are several higher velocities, which appear to be integral multiples of 96 cm/s. The velocities apparently depend on the species, environment, and type of pulse when changed artificially. Measured vertical velocities are larger than horizontal velocities. This may provide a basis for gravitropism and may suggest a cosmic connection. Mean ratios of reciprocals of horizontal and vertical internodal spacings, in simple structure cases, give ratios like 1.25, 1.33, 1.50, 1.66, 3.0, and likely others. These ratios appear to confirm the velocity measurements since ratios of integral multiples of 96 cm/s give the same ratios. The versatility of plant growth is enhanced by the velocity variability.     

Variation in velocity of cytoplasmic streaming and gravity effect in characean internodal cells measured by laser-Doppler-velocimetry

Summary Velocities of cytoplasmic streaming were measured in internodal cells ofNitella flexilis L. andChara corallina Klein ex Willd. by laser-Doppler-velocimetry to investigate the possibility of non-statolith-based perception of gravity. This was recently proposed, based on a report of gravity-dependent polarity of cytoplasmic streaming. Our measurements revealed large spatial and temporal variation in streaming velocity within a cell, independent of the position of the cell with respect to the direction of gravity. In 58% of the horizontally positioned cells the velocities of acropetal and basipetal streaming, measured at opposite locations in the cell, differed significantly. In 45% of these, basipetal streaming was faster than acropetal streaming. In 60% of the vertically positioned cells however the difference was significant, downward streaming was faster in only 61% of these. When cell positions were changed from vertical to horizontal and vice versa the cells reacted variably. A significant difference between velocities in one direction, before and after the change, was observed in approx. 70% of the measurements, but the velocity was faster in the downward direction, as the second position, in only 70% of the significantly different. The ratio of basipetal to acropetal streaming velocities at opposite locations of a cell was quite variable within groups of cells with a particular orientation (horizontal, normal vertical, inverted vertical). On average, however, the ratio was close to 1.00 in the horizontal position and approx. 1.03 in the normal vertical position (basipetal streaming directed downwards), which indicates a small direct effect of gravity on streaming velocity. Individual cells, however, showed an increased, as well as a decreased, ratio when moved from the horizontal to the vertical position. No discernible effect of media (either Ca^{2 +}-buffered medium or 1.2% agar in distilled water) on the streaming velocities was observed. The above mentioned phenomenon of graviperception is not supported by our data.Abbreviations g gravitational acceleration (9.81 m/s²) - LDV laser-Doppler-velocimetry - VR velocity ratio Dedicated to Professor Peter Sitte on the occasion of his 65th birthday     

The role of cutaneous feedback for anticipatory grip force adjustments during object movements and externally imposed variation of the direction of gravity

Grip force adjustments to changes of object loading induced by external changes of the direction of gravity during discrete arm movements with a grasped object were analyzed during normal and anesthetized finger sensibility. Two subjects were seated upright in a rotatable chair and rotated backwards into a horizontal position during discrete movements with a hand-held instrumented object. The movement direction varied from vertical to horizontal inducing corresponding changes in the direction of gravity, but the orientation of the movement in relation to the body remained unaffected. During discrete vertical movements a maximum of load force occurs early in upward and late in downward movements; during horizontal movements two load force peaks result from both acceleratory and deceleratory phases of the movement. During performance with normal finger sensibility grip force was modulated in parallel with fluctuations of load force during vertical and horizontal movements. The grip force profile adopted to the varying load force profile during the transition from the vertical to the horizontal position. The maximum grip force occurred at the same time of maximum load force irrespective of the movement plane. During both subjects' first experience of digital anesthesia the object slipped from the grasp during rotation to the horizontal plane. During the following trials with anesthetized fingers subjects substantially increased their grip forces, resulting in elevated force ratios between maximum grip and load force. However, grip force was still modulated with the movement-induced load fluctuations and maximum grip force coincided with maximum load force during vertical and horizontal movements. This implies that the elevated force ratio between maximum grip and load force does not alter the feedforward system of grip force control. Cutaneous afferent information from the grasping digits seems to be important for the economic scaling of the grip force magnitude according to the actual loading conditions and for reactive grip force adjustments in response to load perturbations. However, it plays a subordinate role for the precise anticipatory temporal coupling between grip and load forces during voluntary object manipulation.     

The role of cutaneous feedback for anticipatory grip force adjustments during object movements and externally imposed variation of the direction of gravity

Grip force adjustments to changes of object loading induced by external changes of the direction of gravity during discrete arm movements with a grasped object were analyzed during normal and anesthetized finger sensibility. Two subjects were seated upright in a rotatable chair and rotated backwards into a horizontal position during discrete movements with a hand-held instrumented object. The movement direction varied from vertical to horizontal inducing corresponding changes in the direction of gravity, but the orientation of the movement in relation to the body remained unaffected. During discrete vertical movements a maximum of load force occurs early in upward and late in downward movements; during horizontal movements two load force peaks result from both acceleratory and deceleratory phases of the movement. During performance with normal finger sensibility grip force was modulated in parallel with fluctuations of load force during vertical and horizontal movements. The grip force profile adopted to the varying load force profile during the transition from the vertical to the horizontal position. The maximum grip force occurred at the same time of maximum load force irrespective of the movement plane. During both subjects' first experience of digital anesthesia the object slipped from the grasp during rotation to the horizontal plane. During the following trials with anesthetized fingers subjects substantially increased their grip forces, resulting in elevated force ratios between maximum grip and load force. However, grip force was still modulated with the movement-induced load fluctuations and maximum grip force coincided with maximum load force during vertical and horizontal movements. This implies that the elevated force ratio between maximum grip and load force does not alter the feedforward system of grip force control. Cutaneous afferent information from the grasping digits seems to be important for the economic scaling of the grip force magnitude according to the actual loading conditions and for reactive grip force adjustments in response to load perturbations. However, it plays a subordinate role for the precise anticipatory temporal coupling between grip and load forces during voluntary object manipulation.     

Kinematics of soft-bodied, legged locomotion in Manduca sexta larvae

Caterpillar crawling is distinct from that of worms and molluscs; it consists of a series of steps in different body segments that can be compared to walking and running in animals with stiff skeletons. Using a three-dimensional kinematic analysis of horizontal crawling in Manduca sexta, the tobacco hornworm, we found that the phase of vertical displacement in the posterior segments substantially led changes in horizontal velocity and the segments appeared to pivot around the attached claspers. Both of the motions occur during vertebrate walking. In contrast, vertical displacement and horizontal velocity in the anterior proleg-bearing segments were in phase, as expected for running gaits coupled by elastic storage. We propose that this kinematic similarity to running results from the muscular compression and release of elastic tissues. As evidence in support of this proposal, the compression and extension of each segment were similar to harmonic oscillations in a spring, although changes in velocity were 70 degrees out of phase with displacement, suggesting that the spring was damped. Measurements of segment length within, and across, intersegmental boundaries show that some of these movements were caused by folding of the body wall between segments. These findings demonstrate that caterpillar crawling is not simply the forward progression of a peristaltic wave but has kinetic components that vary between segments. Although these movements can be compared to legged locomotion in animals with stiff skeletons, the underlying mechanisms of caterpillar propulsion, and in particular the contribution of elastic tissues, remain to be discovered.     

Three-Dimensional Manipulations of Surface Plasmon Polariton Wave Propagation

The plasmonic integrated circuit, a potential application of surface plasmon polaritons (SPPs), can manipulate an SPP wave propagating on a metal surface in a way similar to electronic circuits. Here, we propose the concept of three-dimensional (3D) SPP wave manipulation: control of an SPP wave propagating in both the horizontal direction and the vertical direction. A hole set in the film can guide an SPP wave in the vertical direction. In the horizontal direction, two holographic groove patterns are used to focus an incident SPP wave on one surface of the film to the hole and control the divergent SPP waves transmitted from the hole on the other metal surface, respectively. The holographic groove patterns are designed via the methodology of surface electromagnetic wave holography. 3D finite-difference time-domain method simulations show a good performance of the 3D manipulation via these designed holographic groove patterns.

     

Rotation axes for analysis of gravity effects on plant organs

Epinastic curvatures of branches of Coleus blumei Benth. and the growth pattern of wheat (Triticum aestivum L.) seedlings on clinostats were used for bioassay of rotation methods for preventing growth responses to gravity. Tumbling a plant end over end was found to be just as effective as rotation about its horizontal axis. The results support the reliability of data from experiments in which an entire plant is rotated about a single horizontal line with only part or none of its immature tissues in horizontal orientation to gravity.     

Looking into the water with a facet eye

The apparent velocity and apparent size of objects approaching an animal strongly depend on their size, their position within the visual field and on the path along which they move. It also makes a considerable difference whether both the animal and the objects are in air, or whether the animal looks from one medium into the other, as is the case in animals that live at the water surface. To systematically investigate this situation we calculated apparent velocity and size in the midsagittal plane of approaching objects of flat horizontal, rodlike vertical and spherical shape. We confine our investigation to object movements along straight horizontal and vertical paths and consider the situation for eyes with and without acute zones. The apparent velocity of an approaching object is low in the far field and increases rapidly close to the eye in both air and water. Along horizontal paths and close to the animal it is higher in water than in air. Along vertical paths and close to the animal it is higher in air than in water. Both relationships are exaggerated when there is an acute zone for vertical resolution along the animal's horizon. Boundary curves are calculated along which an approaching object is seen by a linearly increasing number of receptors. The change of apparent size is characterized by the density of these lines. Below the water surface the change of apparent size is similar to that of the apparent velocity. The apparent size of a horizontally elongated object approaching along a horizontal path increases steadily, that of a vertically elongated object along a vertical line initially increases, but can decrease again near the water surface. An acute zone substantially changes the shape and density of these contour lines. The biological consequences of these effects are discussed.     

How do low horizontal forces produce disproportionately high torques in human locomotion?

Although horizontal ground forces are only approximately 15% of vertical forces, they account for 47% and 33% of the metabolic cost in walking and running. To explain these disproportionately high metabolic costs, we hypothesized that low horizontal ground forces generate relatively high torques on body segments during locomotion and this is mediated by long moment arms. We compared external force moment arms and discreet torques applied to the body segments by horizontal and vertical forces during walking and running. Sixteen subjects (21.9+/-1.9 years) walked at 1.5m/s and ten subjects (23.2+/-2.0 years) ran at 3.83 m/s. Segmental torques in the sagittal plane were partitioned into components due to horizontal and vertical forces and quantified by their angular impulses. The mean (+/-S.E.) ratios of horizontal to vertical ground forces (GF ratio) and angular impulses (AI ratio) in walking were 0.131 (+/-0.003, 95% confidence interval (CI) 0.124-0.137) and 0.530 (+/-0.018, CI 0.497-0.569). Results were similar in running. In both gaits the AI ratios were significantly greater than the GF ratios because the respective CI's did not overlap. The horizontal forces produced 53% and 41% as much angular impulse on the body segments, as did the vertical forces in walking and running despite being only 13% as large. In the two movements the moment arms for the horizontal forces averaged across foot, leg, thigh, and trunk body segments were 3.8 fold larger than those for the vertical forces. The data supported the hypothesis and suggest that the relatively low horizontal vs. vertical forces accounted for a disproportionately higher percentage of the angular impulses placed on the body segments and this effect was due to relatively long moment arms for horizontal forces. These results partially explain the relatively large metabolic cost of generating relatively low horizontal forces.     

Clinostating effects on Apogee wheat resistance to wheat streak mosaic virus

The effects of vertical and horizontal clinostating with container velocity 2 rpm and platform velocity 1 rpm on Apogee wheat resistance to wheat streak mosaic virus (WSMV) were studied. For the first time the yield of grain was obtained from both, healthy and the WSMV-infected wheat plants under conditions of simulated microgravity (clinorotation) which is accounted for by the inclusion of various mechanisms in the process of plant adaptation to the factors of stresses. For the first time the stages of viral infection development were elucidated under the conditions of prolonged clinostating and the presence of the WSMV was detected in wheat plants on the 18th day after inoculation employing the method of polymerase chain reaction (PCR). In the test variant with vertical clinostating (R=1,0) the least favorable conditions for viral reproduction were identified. In the same variant the increases of the total nitrogen content in the ears were observed which may be an evidence of interferon-like protein synthesis in plant similar to the interferon system functioning in animals.     

Eye Movements in Ephedrone-Induced Parkinsonism

Patients with ephedrone parkinsonism (EP) show a complex, rapidly progressive, irreversible, and levodopa non-responsive parkinsonian and dystonic syndrome due to manganese intoxication. Eye movements may help to differentiate parkinsonian syndromes providing insights into which brain networks are affected in the underlying disease, but they have never been systematically studied in EP. Horizontal and vertical eye movements were recorded in 28 EP and compared to 21 Parkinson''s disease (PD) patients, and 27 age- and gender-matched healthy subjects using standardized oculomotor tasks with infrared videooculography. EP patients showed slow and hypometric horizontal saccades, an increased occurrence of square wave jerks, long latencies of vertical antisaccades, a high error rate in the horizontal antisaccade task, and made more errors than controls when pro- and antisaccades were mixed. Based on oculomotor performance, a direct differentiation between EP and PD was possible only by the velocity of horizontal saccades. All remaining metrics were similar between both patient groups. EP patients present extensive oculomotor disturbances probably due to manganese-induced damage to the basal ganglia, reflecting their role in oculomotor system.     

Trunk and upper limb muscle activation during flat and topspin forehand drives in young tennis players

This study compared EMG activity of young tennis players' muscles during forehand drives in two groups, GD-those able to raise by more than 150% the vertical velocity of racket-face at impact from flat to topspin forehand drives, and GND, those not able to increase their vertical velocity to the same extent. Upper limb joint angles, racket-face velocities, and average EMGrms values, were studied. At similar joint angles, a fall in horizontal velocity and a rise in racket-face vertical velocity from flat to topspin forehand drives were observed. Shoulder muscle activity rose from flat to topspin forehand drives in GND, but not for drives in GD. Forearm muscle activity reduced from flat to topspin forehand drives in GD, but muscle activation was similar in GND. The results show that radial deviation increased racket-face vertical velocity more at impact from the flat to topspin forehand drives than shoulder abduction.     

Exchange between littoral and pelagic waters in a stratified lake due to wind-induced motions: Lake Kinneret, Israel

We used field data and three-dimensional numerical simulations to investigate the exchange processes between the lake boundaries (littoral) and lake interior (pelagic) due to wind-induced motions in Lake Kinneret, Israel. The field data showed large-scale metalimnion oscillations with amplitudes up to 10 m in response to westerly diurnal winds, the existence of a well-defined turbidity intrusion into the metalimnion of the lake and a benthic boundary layer (BBL). The observed vertical and horizontal movements of the turbidity were explained by the vertical and horizontal advection associated with basin-scale wave motions and a residual circulation set up by the basin-scale motions and the wind field. The horizontal advective transport in the metalimnion, associated with the velocities induced by the basin-scale mode-two Poincaré wave, provided the primary control for the exchange between the lake boundaries and lake interior on daily time scales. Detailed comparison of simulation results with both temperature and velocity profiles revealed excellent agreement for time scales from hours to days. The validated numerical model was used to extract the residual motions that provided the secondary exchange flows, in the various regions of the lake on time scales from days to weeks. The residual motion in the surface layer consisted of a combination of an anticlockwise gyre in the western half of the lake and a weak clockwise gyre along the eastern boundary that were forced by the curl of the wind field. In the metalimnion, residual motions were predominantly forced by the basin-scale internal wave motions and an anticlockwise gyre established itself throughout the whole basin. Lastly in the BBL, residual motions consisted of an anticlockwise gyre, but the geographic distribution was patchy. Together these results provide new insight into a proper characterization of the processes underlying the flux paths of water and particles between the lake boundaries and lake interior. Handling editor: L. Naselli-Flores     

Effects of gravity on positive phototaxis in fruit fly Drosophila melanogaster

Geotaxis and phototaxis are movements in response to gravity and light, respectively, and are commonly observed in nature. The interactions between these two types of movement have been shown to confer ecological advantages to many taxa. Although several studies have been conducted on phototaxis and geotaxis in various organisms, reports on the interactions between positive phototaxis and negative geotaxis are lacking. In the fruit fly, Drosophila melanogaster, any direct interactions that exist between positive phototaxis and negative geotaxis are yet to be determined and the ecological significance of such interactions remains unclear. In the present study, the effects of gravity on positive phototaxis in a Y‐maze were investigated using the Canton‐S wild type and gravity‐sensing‐deficient pyx³ mutant fruit flies. Gravity sensing was not necessary for horizontal positive phototaxis, but was required for vertical positive phototaxis. These results suggest that gravitoreception may selectively modulate positive phototaxis depending on the vertical and horizontal movements of the fruit flies.     

Within-plant distribution of cotton aphids, Aphis gossypii Glover (Hemiptera: Aphididae), in Bt and non-Bt cotton fields

Knowledge of the vertical and horizontal distribution of Aphis gossypii Glover (Hemiptera: Aphididae) on genetically modified cotton plants over time could help optimize decision-making in integrated cotton aphid management programs. Therefore, the aim of the present study was to determine the vertical and horizontal distribution of A. gossypii in non-transgenic Bt cotton and transgenic Bt-cotton over time during two cotton seasons by examining plants throughout the seasons. There was no significant interaction between years and cotton cultivar treatments for apterous or alate aphids. Considering year-to-year data, analyses on season-long averages of apterous or alate aphids showed that aphid densities per plant did not differ among years. The number of apterous aphids found per plant for the Bt transgenic cultivar (2427 apterous aphids per plant) was lower than for its isoline (3335 apterous aphids per plant). The number of alate aphids found per plant on the Bt transgenic cultivar (12.28 alate aphids per plant) was lower than for the isoline (140.56 alate aphids per plant). With regard to the vertical distribution of apterous aphids or alate aphids, there were interactions between cotton cultivar, plant age and plant region. We conclude that in comparison to non-Bt cotton (DP 4049), Bt cotton (DP 404 BG (Bollgard)) has significant effects on the vertical, horizontal, spatial and temporal distribution patterns of A. gossypii, showing changes in its distribution behaviour inside the plant as the cotton crop develops. The results of our study are relevant for understanding the vertical and horizontal distribution of A. gossypii on Bt cotton cultivar (DP 404 BG (Bollgard)) and on its isoline (DP 4049), and could be useful in decision-making, implementing controls and determining the timing of population peaks of this insect.     

An oscillation theory of handwriting

Handwriting production is viewed as a constrained modulation of an underlying oscillatory process. Coupled oscillations in horizontal and vertical directions produce letter forms, and when superimposed on a rightward constant velocity horizontal sweep result in spatially separated letters. Modulation of the vertical oscillation is responsible for control of letter height. Modulation of the horizontal oscillation is responsible for control of corner shape through altering phase or amplitude. The vertical velocity zero crossing in the velocity space diagram is important from the standpoint of control. Changing the horizontal velocity value at this zero crossing controls corner shape. Changing the slope at this zero crossing controls writing slant. The corner shape and slant constraints completely determine the amplitude and phase relations between the two oscillations. This theory applies generally to a number of acceleration oscillation patterns such as sinusoidal rectangular and trapezoidal oscillations. The oscillation theory also provides an explanation for how handwriting might degenerate with speed. An implementation of the theory in the context of the spring muscle model is developed. Here sinusoidal oscillations arise from a purely mechanical source; orthogonal antagonistic spring pairs generate particular cycloids depending on the initial conditions. Modulating between cycloids can be achieved by changing the spring zero settings at the appropriate times. Frequency can be modulated either by shifting between coactivation and alternating activation of the antagonistic springs or by presuming variable spring constant springs. An acceleration and position measuring apparatus was developed for measurements of human handwriting. Measurements of human writing are consistent with the oscillation theory.     

Growth and development of Potamogeton distinctus in an irrigation pond in SW Japan

Growth and development of a population of Potamogeton distinctus were studied over one vegetation period.
The morphological structure of the species is described. Maximum development of the size of shoot complexes occurred in August (up to 12 g dry weight/shoot complex). The plant exhibits a regular growth cycle with continuous horizontal and vertical growth that is illustrated by the development of horizontal shoots, vertical shoots and different plant parts. The lower horizontal shoots allow an extension up to 1.8 m from the point of germination. Before the rainy season in June the 1st to 5th generations of vertical shoots mainly contribute to the standing crop of the plants' stand, and after the rainy season the 5th to 9th generation shoots are predominant. Floating leaves make up the main proportion of biomass during the whole growth cycle. Almost 100% of the shoot complexes were found flowering and producing numerous seeds. Up to 16% of the standing crop are allocated to inflorescence and seeds in August. Turions failed to form that year because of rapid desiccation of the habitat. The observed average lifetime of a vertical shoot is appr. 65 days, the estimated turnover rate is between 2.15 and 2.27.     

THE EVOLUTION OF VIRULENCE IN PATHOGENS WITH VERTICAL AND HORIZONTAL TRANSMISSION

The idea that vertical transmission of parasites selects for lower virulence is widely accepted. However, little theoretical work has considered the evolution of virulence for parasites with mixed horizontal plus vertical transmission. Many human, animal, and plant parasites are transmitted both vertically and horizontally, and some horizontal transmission is generally necessary to maintain parasites at all. We present a population-dynamical model for the evolution of virulence when both vertical and horizontal transmission are present. In the simplest such model, up to two infectious strains can coexist within one host population. Virulent, vertically transmitted pathogens can persist in a population when they provide protection against more virulent, horizontally transmitted strains. When virulence is maintained by a correlation with horizontal transmission rates, increased levels of vertical transmission always lower the evolutionarily stable (ESS) level of virulence. Contrary to existing theory, however, increases in opportunities for horizontal transmission also lower the ESS level of virulence. We explain these findings in light of earlier work and confirm them in simulations including imperfect vertical transmission. We describe further simulations, in which both vertical and horizontal transmission rates are allowed to evolve. The outcome of these simulations depends on whether high levels of vertical transmission are possible with low virulence. Finally, we argue against the notion of a virulence-avirulence continuum between horizontal and vertical transmission, and discuss our results in relation to empirical studies of transmission and virulence.     

Horizontal and vertical transmission of viruses in the honey bee, Apis mellifera

The most crucial stage in the dynamics of virus infections is the mode of virus transmission. In general, transmission of viruses can occur through two pathways: horizontal and vertical transmission. In horizontal transmission, viruses are transmitted among individuals of the same generation, while vertical transmission occurs from mothers to their offspring. Because of its highly organized social structure and crowded population density, the honey bee colony represents a risky environment for the spread of disease infection. Like other plant and animal viruses, bee viruses use different survival strategies, including utilization of both horizontal and vertical routes, to transmit and maintain levels in a host population. In this review, we explore the current knowledge about the honey bee viruses and transmission routes of bee viruses. In addition, different transmission strategies on the persistence and dynamics of host-pathogen interactions are also discussed.