Eye-Hand Coordination during Visuomotor Adaptation with Different Rotation Angles: Effects of Terminal Visual Feedback

This study examined adaptive changes of eye-hand coordination during a visuomotor rotation task under the use of terminal visual feedback. Young adults made reaching movements to targets on a digitizer while looking at targets on a monitor where the rotated feedback (a cursor) of hand movements appeared after each movement. Three rotation angles (30°, 75° and 150°) were examined in three groups in order to vary the task difficulty. The results showed that the 30° group gradually reduced direction errors of reaching with practice and adapted well to the visuomotor rotation. The 75° group made large direction errors of reaching, and the 150° group applied a 180° reversal shift from early practice. The 75°and 150° groups, however, overcompensated the respective rotations at the end of practice. Despite these group differences in adaptive changes of reaching, all groups gradually adapted gaze directions prior to reaching from the target area to the areas related to the final positions of reaching during the course of practice. The adaptive changes of both hand and eye movements in all groups mainly reflected adjustments of movement directions based on explicit knowledge of the applied rotation acquired through practice. Only the 30° group showed small implicit adaptation in both effectors. The results suggest that by adapting gaze directions from the target to the final position of reaching based on explicit knowledge of the visuomotor rotation, the oculomotor system supports the limb-motor system to make precise preplanned adjustments of reaching directions during learning of visuomotor rotation under terminal visual feedback.     

Simultaneous Processing of Information on Multiple Errors in Visuomotor Learning

The proper association between planned and executed movements is crucial for motor learning because the discrepancies between them drive such learning. Our study explored how this association was determined when a single action caused the movements of multiple visual objects. Participants reached toward a target by moving a cursor, which represented the right hand’s position. Once every five to six normal trials, we interleaved either of two kinds of visual perturbation trials: rotation of the cursor by a certain amount (±15°, ±30°, and ±45°) around the starting position (single-cursor condition) or rotation of two cursors by different angles (+15° and −45°, 0° and 30°, etc.) that were presented simultaneously (double-cursor condition). We evaluated the aftereffects of each condition in the subsequent trial. The error sensitivity (ratio of the aftereffect to the imposed visual rotation) in the single-cursor trials decayed with the amount of rotation, indicating that the motor learning system relied to a greater extent on smaller errors. In the double-cursor trials, we obtained a coefficient that represented the degree to which each of the visual rotations contributed to the aftereffects based on the assumption that the observed aftereffects were a result of the weighted summation of the influences of the imposed visual rotations. The decaying pattern according to the amount of rotation was maintained in the coefficient of each imposed visual rotation in the double-cursor trials, but the value was reduced to approximately 40% of the corresponding error sensitivity in the single-cursor trials. We also found a further reduction of the coefficients when three distinct cursors were presented (e.g., −15°, 15°, and 30°). These results indicated that the motor learning system utilized multiple sources of visual error information simultaneously to correct subsequent movement and that a certain averaging mechanism might be at work in the utilization process.     

Spatial Interactions between Successive Eye and Arm Movements: Signal Type Matters

Spatial interactions between consecutive movements are often attributed to inhibition of return (IOR), a phenomenon in which responses to previously signalled locations are slower than responses to unsignalled locations. In two experiments using peripheral target signals offset by 0°, 90°, or 180°, we show that consecutive saccadic (Experiment 1) and reaching (Experiment 3) responses exhibit a monotonic pattern of reaction times consistent with the currently established spatial distribution of IOR. In contrast, in two experiments with central target signals (i.e., arrowheads pointing at target locations), we find a non-monotonic pattern of reaction times for saccades (Experiment 2) and reaching movements (Experiment 4). The difference in the patterns of results observed demonstrates different behavioral effects that depend on signal type. The pattern of results observed for central stimuli are consistent with a model in which neural adaptation is occurring within motor networks encoding movement direction in a distributed manner.     

Specificity of sensorimotor learning and the neural code: Neuronal representations in the primary motor cortex

Human studies show that the learning of a new sensorimotor mapping that requires adaptation to directional errors is local and generalizes poorly to untrained directions. We trained monkeys to learn new visuomotor rotations for only one target in space and recorded neuronal activity in the primary motor cortex before, during and after learning. Similar to humans, the monkeys showed poor transfer of learning to other directions, as observed by behavioral aftereffects for untrained directions. To test for internal representations underlying these changes, we compared two features of neuronal activity before and after learning: changes in firing rates and changes in information content. Specific elevations of firing rate were only observed in a subpopulation of cells in the motor cortex with directional properties corresponding to the locally learned rotation; namely cells only showed plasticity if their preferred direction was near the training one. We applied measures from information theory to probe for learning-related changes in the neuronal code. Single cells conveyed more information about the direction of movement and this specific improvement in encoding was correlated with an increase in the slope of the neurons' tuning curve. Further, the improved information after learning enabled a more accurate reconstruction of movement direction from neuronal populations. Our findings suggest a neural mechanism for the confined generalization of a newly acquired internal model by showing a tight relationship between the locality of learning and the properties of neurons. They also provide direct evidence for improvement in the neural code as a result of learning.     

Rethinking motor learning and savings in adaptation paradigms: model-free memory for successful actions combines with internal models

Although motor learning is likely to involve multiple processes, phenomena observed in error-based motor learning paradigms tend to be conceptualized in terms of only a single process: adaptation, which occurs through updating an internal model. Here we argue that fundamental phenomena like movement direction biases, savings (faster relearning), and interference do not relate to adaptation but instead are attributable to two additional learning processes that can be characterized as model-free: use-dependent plasticity and operant reinforcement. Although usually "hidden" behind adaptation, we demonstrate, with modified visuomotor rotation paradigms, that these distinct model-based and model-free processes combine to learn an error-based motor task. (1) Adaptation of an internal model channels movements toward successful error reduction in visual space. (2) Repetition of the newly adapted movement induces directional biases toward the?repeated movement. (3) Operant reinforcement through association of the adapted movement with successful error reduction is responsible for savings.     

Coordinate Representations for Interference Reduction in Motor Learning

When opposing force fields are presented alternately or randomly across trials for identical reaching movements, subjects learn neither force field, a behavior termed ‘interference’. Studies have shown that a small difference in the endpoint posture of the limb reduces this interference. However, any difference in the limb’s endpoint location typically changes the hand position, joint angles and the hand orientation making it ambiguous as to which of these changes underlies the ability to learn dynamics that normally interfere. Here we examine the extent to which each of these three possible coordinate systems—Cartesian hand position, shoulder and elbow joint angles, or hand orientation—underlies the reduction in interference. Subjects performed goal-directed reaching movements in five different limb configurations designed so that different pairs of these configurations involved a change in only one coordinate system. By specifically assigning clockwise and counter-clockwise force fields to the configurations we could create three different conditions in which the direction of the force field could only be uniquely distinguished in one of the three coordinate systems. We examined the ability to learn the two fields based on each of the coordinate systems. The largest reduction of interference was observed when the field direction was linked to the hand orientation with smaller reductions in the other two conditions. This result demonstrates that the strongest reduction in interference occurred with changes in the hand orientation, suggesting that hand orientation may have a privileged role in reducing motor interference for changes in the endpoint posture of the limb.     

Directional Coordination of Thumb and Finger Forces during Precision Pinch

The human opposable thumb enables the hand to perform dexterous manipulation of objects, which requires well-coordinated digit force vectors. This study investigated the directional coordination of force vectors generated by the thumb and index finger during precision pinch. Fourteen right-handed, healthy subjects were instructed to exert pinch force on an externally stabilized apparatus with the pulps of the thumb and index finger. Subjects applied forces to follow a force-ramp profile that linearly increased from 0 to 12 N and then decreased to 0 N, at a rate of ±3 N/s. Directional relationships between the thumb and index finger force vectors were quantified using the coordination angle (CA) between the force vectors. Individual force vectors were further analyzed according to their projection angles (PAs) with respect to the pinch surface planes and the shear angles (SAs) within those planes. Results demonstrated that fingertip force directions were dependent on pinch force magnitude, especially at forces below 2 N. Hysteresis was observed in the force-CA relationship for increasing and decreasing forces and fitted with exponential models. The fitted asymptotic values were 156.0±6.6° and 150.8±9.3° for increasing and decreasing force ramps, respectively. The PA of the thumb force vector deviated further from the direction perpendicular to the pinching surface planes than that of the index finger. The SA showed that the index finger force vector deviated in the ulnar-proximal direction, whereas the thumb switched its force between the ulnar-proximal and radial-proximal directions. The findings shed light on the effects of anatomical composition, biomechanical function, and neuromuscular control in coordinating digit forces during precision pinch, and provided insight into the magnitude-dependent force directional control which potentially affects a range of dexterous manipulations.     

Trunk Muscle Activation at the Initiation and Braking of Bilateral Shoulder Flexion Movements of Different Amplitudes

The aim of this study was to investigate if trunk muscle activation patterns during rapid bilateral shoulder flexions are affected by movement amplitude. Eleven healthy males performed shoulder flexion movements starting from a position with arms along sides (0°) to either 45°, 90° or 180°. EMG was measured bilaterally from transversus abdominis (TrA), obliquus internus (OI) with intra-muscular electrodes, and from rectus abdominis (RA), erector spinae (ES) and deltoideus with surface electrodes. 3D kinematics was recorded and inverse dynamics was used to calculate the reactive linear forces and torque about the shoulders and the linear and angular impulses. The sequencing of trunk muscle onsets at the initiation of arm movements was the same across movement amplitudes with ES as the first muscle activated, followed by TrA, RA and OI. All arm movements induced a flexion angular impulse about the shoulders during acceleration that was reversed during deceleration. Increased movement amplitude led to shortened onset latencies of the abdominal muscles and increased level of activation in TrA and ES. The activation magnitude of TrA was similar in acceleration and deceleration where the other muscles were specific to acceleration or deceleration. The findings show that arm movements need to be standardized when used as a method to evaluate trunk muscle activation patterns and that inclusion of the deceleration of the arms in the analysis allow the study of the relationship between trunk muscle activation and direction of perturbing torque during one and the same arm movement.     

Learning-induced improvement in encoding and decoding of specific movement directions by neurons in the primary motor cortex

Many recent studies describe learning-related changes in sensory and motor areas, but few have directly probed for improvement in neuronal coding after learning. We used information theory to analyze single-cell activity from the primary motor cortex of monkeys, before and after learning a local rotational visuomotor task. We show that after learning, neurons in the primary motor cortex conveyed more information about the direction of movement and did so with relation to their directional sensitivity. Similar to recent findings in sensory systems, this specific improvement in encoding is correlated with an increase in the slope of the neurons' tuning curve. We further demonstrate that the improved information after learning enables a more accurate reconstruction of movement direction from neuronal populations. Our results suggest that similar mechanisms govern learning in sensory and motor areas and provide further evidence for a tight relationship between the locality of learning and the properties of neurons; namely, cells only show plasticity if their preferred direction is near the training one. The results also suggest that simple learning tasks can enhance the performance of brain–machine interfaces.     

Avian magnetic compass can be tuned to anomalously low magnetic intensities

The avian magnetic compass works in a fairly narrow functional window around the intensity of the local geomagnetic field, but adjusts to intensities outside this range when birds experience these new intensities for a certain time. In the past, the geomagnetic field has often been much weaker than at present. To find out whether birds can obtain directional information from a weak magnetic field, we studied spontaneous orientation preferences of migratory robins in a 4 µT field (i.e. a field of less than 10 per cent of the local intensity of 47 µT). Birds can adjust to this low intensity: they turned out to be disoriented under 4 µT after a pre-exposure time of 8 h to 4 µT, but were able to orient in this field after a total exposure time of 17 h. This demonstrates a considerable plasticity of the avian magnetic compass. Orientation in the 4 µT field was not affected by local anaesthesia of the upper beak, but was disrupted by a radiofrequency magnetic field of 1.315 MHz, 480 nT, suggesting that a radical-pair mechanism still provides the directional information in the low magnetic field. This is in agreement with the idea that the avian magnetic compass may have developed already in the Mesozoic in the common ancestor of modern birds.     

Repetitive Transcranial Magnetic Stimulation Attenuates the Perception of Force Output Production in Non-Exercised Hand Muscles after Unilateral Exercise

We examined whether unilateral exercise creates perception bias in the non-exercised limb and ascertained whether rTMS applied to the primary motor cortex (M1) interferes with this perception. All participants completed 4 interventions: 1) 15-min learning period of intermittent isometric contractions at 35% MVC with the trained hand (EX), 2) 15-min learning period of intermittent isometric contractions at 35% MVC with the trained hand whilst receiving rTMS over the contralateral M1 (rTMS+EX); 3) 15-min of rTMS over the ‘trained’ M1 (rTMS) and 4) 15-min rest (Rest). Pre and post-interventions, the error of force output production, the perception of effort (RPE), motor evoked potentials (MEPs) and compound muscle action potentials (CMAPs) were measured in both hands. EX did not alter the error of force output production in the trained hand (Δ3%; P>0.05); however, the error of force output production was reduced in the untrained hand (Δ12%; P<0.05). rTMS+EX and rTMS alone did not show an attenuation in the error of force output production in either hand. EX increased RPE in the trained hand (9.1±0.5 vs. 11.3±0.7; P<0.01) but not the untrained hand (8.8±0.6 vs. 9.2±0.6; P>0.05). RPE was significantly higher after rTMS+EX in the trained hand (9.2±0.5 vs. 10.7±0.7; P<0.01) but ratings were unchanged in the untrained hand (8.5±0.6 vs. 9.2±0.5; P>0.05). The novel finding was that exercise alone reduced the error in force output production by over a third in the untrained hand. Further, when exercise was combined with rTMS the transfer of force perception was attenuated. These data suggest that the contralateral M1 of the trained hand might, in part, play an essential role for the transfer of force perception to the untrained hand.     

Hawk Eyes I: Diurnal Raptors Differ in Visual Fields and Degree of Eye Movement

Background

Different strategies to search and detect prey may place specific demands on sensory modalities. We studied visual field configuration, degree of eye movement, and orbit orientation in three diurnal raptors belonging to the Accipitridae and Falconidae families.

Methodology/Principal Findings

We used an ophthalmoscopic reflex technique and an integrated 3D digitizer system. We found inter-specific variation in visual field configuration and degree of eye movement, but not in orbit orientation. Red-tailed Hawks have relatively small binocular areas (∼33°) and wide blind areas (∼82°), but intermediate degree of eye movement (∼5°), which underscores the importance of lateral vision rather than binocular vision to scan for distant prey in open areas. Cooper''s Hawks'' have relatively wide binocular fields (∼36°), small blind areas (∼60°), and high degree of eye movement (∼8°), which may increase visual coverage and enhance prey detection in closed habitats. Additionally, we found that Cooper''s Hawks can visually inspect the items held in the tip of the bill, which may facilitate food handling. American Kestrels have intermediate-sized binocular and lateral areas that may be used in prey detection at different distances through stereopsis and motion parallax; whereas the low degree eye movement (∼1°) may help stabilize the image when hovering above prey before an attack.

Conclusions

We conclude that: (a) there are between-species differences in visual field configuration in these diurnal raptors; (b) these differences are consistent with prey searching strategies and degree of visual obstruction in the environment (e.g., open and closed habitats); (c) variations in the degree of eye movement between species appear associated with foraging strategies; and (d) the size of the binocular and blind areas in hawks can vary substantially due to eye movements. Inter-specific variation in visual fields and eye movements can influence behavioral strategies to visually search for and track prey while perching.     

Covert Anti-Compensatory Quick Eye Movements during Head Impulses

Background

Catch-up saccades during passive head movements, which compensate for a deficient vestibulo-ocular reflex (VOR), are a well-known phenomenon. These quick eye movements are directed toward the target in the opposite direction of the head movement. Recently, quick eye movements in the direction of the head movement (covert anti-compensatory quick eye movements, CAQEM) were observed in older individuals. Here, we characterize these quick eye movements, their pathophysiology, and clinical relevance during head impulse testing (HIT).

Methods

Video head impulse test data from 266 patients of a tertiary vertigo center were retrospectively analyzed. Forty-three of these patients had been diagnosed with vestibular migraine, and 35 with Menière’s disease.

Results

CAQEM occurred in 38% of the patients. The mean CAQEM occurrence rate (per HIT trial) was 11±10% (mean±SD). Latency was 83±30 ms. CAQEM followed the saccade main sequence characteristics and were compensated by catch-up saccades in the opposite direction. Compensatory saccades did not lead to more false pathological clinical head impulse test assessments (specificity with CAQEM: 87%, and without: 85%). CAQEM on one side were associated with a lower VOR gain on the contralateral side (p<0.004) and helped distinguish Menière’s disease from vestibular migraine (p = 0.01).

Conclusion

CAQEM are a common phenomenon, most likely caused by a saccadic/quick phase mechanism due to gain asymmetries. They could help differentiate two of the most common causes of recurrent vertigo: vestibular migraine and Menière’s disease.     

Generalization of motor learning depends on the history of prior action

Generalization of motor learning refers to our ability to apply what has been learned in one context to other contexts. When generalization is beneficial, it is termed transfer, and when it is detrimental, it is termed interference. Insight into the mechanism of generalization may be acquired from understanding why training transfers in some contexts but not others. However, identifying relevant contextual cues has proven surprisingly difficult, perhaps because the search has mainly been for cues that are explicit. We hypothesized instead that a relevant contextual cue is an implicit memory of action with a particular body part. To test this hypothesis we considered a task in which participants learned to control motion of a cursor under visuomotor rotation in two contexts: by moving their hand through motion of their shoulder and elbow, or through motion of their wrist. Use of these contextual cues led to three observations: First, in naive participants, learning in the wrist context was much faster than in the arm context. Second, generalization was asymmetric so that arm training benefited subsequent wrist training, but not vice versa. Third, in people who had prior wrist training, generalization from the arm to the wrist was blocked. That is, prior wrist training appeared to prevent both the interference and transfer that subsequent arm training should have caused. To explain the data, we posited that the learner collected statistics of contextual history: all upper arm movements also move the hand, but occasionally we move our hands without moving the upper arm. In a Bayesian framework, history of limb segment use strongly affects parameter uncertainty, which is a measure of the covariance of the contextual cues. This simple Bayesian prior dictated a generalization pattern that largely reproduced all three findings. For motor learning, generalization depends on context, which is determined by the statistics of how we have previously used the various parts of our limbs.     

Cognitive Loading Affects Motor Awareness and Movement Kinematics but Not Locomotor Trajectories during Goal-Directed Walking in a Virtual Reality Environment

The primary purpose of this study was to investigate the effects of cognitive loading on movement kinematics and trajectory formation during goal-directed walking in a virtual reality (VR) environment. The secondary objective was to measure how participants corrected their trajectories for perturbed feedback and how participants'' awareness of such perturbations changed under cognitive loading. We asked 14 healthy young adults to walk towards four different target locations in a VR environment while their movements were tracked and played back in real-time on a large projection screen. In 75% of all trials we introduced angular deviations of ±5° to ±30° between the veridical walking trajectory and the visual feedback. Participants performed a second experimental block under cognitive load (serial-7 subtraction, counter-balanced across participants). We measured walking kinematics (joint-angles, velocity profiles) and motor performance (end-point-compensation, trajectory-deviations). Motor awareness was determined by asking participants to rate the veracity of the feedback after every trial. In-line with previous findings in natural settings, participants displayed stereotypical walking trajectories in a VR environment. Our results extend these findings as they demonstrate that taxing cognitive resources did not affect trajectory formation and deviations although it interfered with the participants'' movement kinematics, in particular walking velocity. Additionally, we report that motor awareness was selectively impaired by the secondary task in trials with high perceptual uncertainty. Compared with data on eye and arm movements our findings lend support to the hypothesis that the central nervous system (CNS) uses common mechanisms to govern goal-directed movements, including locomotion. We discuss our results with respect to the use of VR methods in gait control and rehabilitation.     

Measuring generalization of visuomotor perturbations in wrist movements using mobile phones

Recent studies in motor control have shown that visuomotor rotations for reaching have narrow generalization functions: what we learn during movements in one direction only affects subsequent movements into close directions. Here we wanted to measure the generalization functions for wrist movement. To do so we had 7 subjects performing an experiment holding a mobile phone in their dominant hand. The mobile phone's built in acceleration sensor provided a convenient way to measure wrist movements and to run the behavioral protocol. Subjects moved a cursor on the screen by tilting the phone. Movements on the screen toward the training target were rotated and we then measured how learning of the rotation in the training direction affected subsequent movements in other directions. We find that generalization is local and similar to generalization patterns of visuomotor rotation for reaching.     

Factors of Force Potentiation Induced by Stretch-Shortening Cycle in Plantarflexors

Muscle force is potentiated by countermovement; this phenomenon is called stretch-shortening cycle (SSC) effect. In this study, we examined the factors strongly related to SSC effect in vivo, focusing on tendon elongation, preactivation, and residual force enhancement. Twelve healthy men participated in this study. Ankle joint angle was passively moved by a dynamometer, with a range of motion from 15° dorsiflexion (DF) to 15° plantarflexion (PF). Muscle contraction was evoked by electrical stimulation, with stimulation timing adjusted to elicit three types of contraction: (1) concentric contraction without preliminary contraction (CON), (2) concentric contraction after preliminary eccentric contraction (ECC), and (3) concentric contraction after preliminary isometric contraction (ISO). Joint torque was recorded at DF5°, PF0°, and PF5°, respectively. SSC effect was calculated as the ratio of joint torque obtained in ECC or ISO with respect to that obtained in CON at the aforementioned three joint angles. SSC effect was prominent in the first half of movement in both ECC (DF5°, 329.3 ± 101.2%; PF0°, 159.2 ± 29.4%; PF5°, 125.5 ± 20.8%) and ISO (DF5°, 276.4 ± 87.0%; PF0°, 134.5 ± 24.5%; PF5°, 106.8 ± 18.0%) conditions. SSC effect was significantly larger in ECC than in ISO at all joint angles (P < 0.001). Even without preliminary eccentric contraction (i.e., ISO condition), SSC effect was clearly large, indicating that a significant part of SSC effect is derived from preactivation. However, the active lengthening-induced force potentiation mechanism (residual force enhancement) also contributes to SSC effect.     

Eye-Head Coordination for Visual Cognitive Processing

We investigated coordinated movements between the eyes and head (“eye-head coordination”) in relation to vision for action. Several studies have measured eye and head movements during a single gaze shift, focusing on the mechanisms of motor control during eye-head coordination. However, in everyday life, gaze shifts occur sequentially and are accompanied by movements of the head and body. Under such conditions, visual cognitive processing influences eye movements and might also influence eye-head coordination because sequential gaze shifts include cycles of visual processing (fixation) and data acquisition (gaze shifts). In the present study, we examined how the eyes and head move in coordination during visual search in a large visual field. Subjects moved their eyes, head, and body without restriction inside a 360° visual display system. We found patterns of eye-head coordination that differed those observed in single gaze-shift studies. First, we frequently observed multiple saccades during one continuous head movement, and the contribution of head movement to gaze shifts increased as the number of saccades increased. This relationship between head movements and sequential gaze shifts suggests eye-head coordination over several saccade-fixation sequences; this could be related to cognitive processing because saccade-fixation cycles are the result of visual cognitive processing. Second, distribution bias of eye position during gaze fixation was highly correlated with head orientation. The distribution peak of eye position was biased in the same direction as head orientation. This influence of head orientation suggests that eye-head coordination is involved in gaze fixation, when the visual system processes retinal information. This further supports the role of eye-head coordination in visual cognitive processing.     

Fluctuation Analysis of Centrosomes Reveals a Cortical Function of Kinesin-1

The actin and microtubule networks form the dynamic cytoskeleton. Network dynamics is driven by molecular motors applying force onto the networks and the interactions between the networks. Here we assay the dynamics of centrosomes in the scale of seconds as a proxy for the movement of microtubule asters. With this assay we want to detect the role of specific motors and of network interaction. During interphase of syncytial embryos of Drosophila, cortical actin and the microtubule network depend on each other. Centrosomes induce cortical actin to form caps, whereas F-actin anchors microtubules to the cortex. In addition, lateral interactions between microtubule asters are assumed to be important for regular spatial organization of the syncytial embryo. The functional interaction between the microtubule asters and cortical actin has been largely analyzed in a static manner, so far. We recorded the movement of centrosomes at 1 Hz and analyzed their fluctuations for two processes—pair separation and individual movement. We found that F-actin is required for directional movements during initial centrosome pair separation, because separation proceeds in a diffusive manner in latrunculin-injected embryos. For assaying individual movement, we established a fluctuation parameter as the deviation from temporally and spatially slowly varying drift movements. By analysis of mutant and drug-injected embryos, we found that the fluctuations were suppressed by both cortical actin and microtubules. Surprisingly, the microtubule motor Kinesin-1 also suppressed fluctuations to a similar degree as F-actin. Kinesin-1 may mediate linkage of the microtubule (+)-ends to the actin cortex. Consistent with this model is our finding that Kinesin-1-GFP accumulates at the cortical actin caps.     

Pollination adaptations of group-by-group stamen movement in a meadow plant with temporal floral closure

Floral sexual organ (stamen and pistil) movements are selective adaptations that have different functions in male-female reproduction and the evolution of flowering plants. However, the significance of stamen movements in the spatial–temporal function and separation of male and female organs has not been experimentally determined in species exhibiting floral temporal closure. The current study investigated the role of slow stamen (group-by-group) movement in male-female sexual function, and the effect of stamen movement on pollen removal, male-male and male-female interference, and mating patterns of Geranium pratense, a plant with temporal floral closure. This species uses stamen group-by-group movement and therefore anther-stigma spatial–temporal separation. Spatial separation (two whorls of stamen and pistil length) was shown to be stronger than temporal separation. We found that stamen movements to the center of the flower increase pollen removal, and the most common pollinators visited more frequently and for longer durations during the male floral stage than during the female floral stage. Petal movements increased both self-pollen deposition rate and sexual interference in G. pratense. The fruit and seed set of naturally and outcrossed pollinated flowers were more prolific than those of self-pollinated flowers. Group-by-group stamen movement, dehiscence of stamens, pistil movement, and male-female spatial–temporal functional separation of G. pratense before floral temporal closure may prevent male-female and stamen-stamen interference and pollen discounting, and may increase pollen removal and cross-pollination.