Distinct adaptation processes underlie multidigit force coordination for dexterous manipulation.

The human sensorimotor system can adapt to various changes in the environmental dynamics by updating motor commands to improve performance after repeated exposure to the same task. However, the characteristics and mechanisms of the adaptation process remain unknown for dexterous manipulation, a unique motor task in which the body physically interacts with the environment with multiple effectors, i.e., digits, in parallel. We addressed this gap by using robotic manipulanda to investigate the changes in the digit force coordination following mechanical perturbation of an object held by tripod grasps. As the participants gradually adapted to lifting the object under perturbations, we quantified two components of digit force coordination. One is the direction-specific manipulation moment that directly counteracts the perturbation, whereas the other one is the direction-independent internal moment that supports the stability and stiffness of the grasp. We found that trial-to-trial improvement of task performance was associated with increased manipulation moment and a gradual decrease of the internal moment. These two moments were characterized by different rates of adaptation. We also examined how these two force coordination components respond to changes in perturbation directions. Importantly, we found that the manipulation moment was sensitive to the extent of repetitive exposure to the previous context that has an opposite perturbation direction, whereas the internal moment did not. However, the internal moment was sensitive to whether the postchange perturbation direction was previously experienced. Our results reveal, for the first time, that two distinct processes underlie the adaptation of multidigit force coordination for dexterous manipulation.NEW & NOTEWORTHY Changes in digit force coordination in multidigit object manipulation were quantified with a novel experimental design in which human participants adapted to mechanical perturbations applied to the object. Our results show that the adaptation of digit force coordination can be characterized by two distinct components that operate at different timescales. We further show that these two components respond to changes in perturbation direction differently.

Molecular mapping for fruit-related traits, and joint identification of candidate genes and selective sweeps for seed size in melon.

Melon is a popular fruit vegetable crop worldwide with diverse morphological variation. We report a high-density genetic map of melon and nine major QTLs with physical region ranging from 43.47 kb to 1.89 Mb. Importantly, two seed-related trait QTLs were repeatedly detected in two environments, and the mapping region was narrowed to 522 kb according to a regional linkage analysis. A total of 40 annotated genes were screened for nonsynonymous variations, of which EVM0009818, involved in cytokinin-activated signaling, was differentially expressed in the young fruits of parents based on RNA-seq. Selective sweep analysis identified 152 sweep signals for seed size, including the two seed-related QTLs and nine homologs that have been verified to regulate seed size in Arabidopsis or rice. This work illustrates the power of a joint analysis combining resequencing-based genetic map for QTL mapping and a combination of KASP genotyping and RNA-seq analysis to facilitate QTL fine mapping.

Fine-mapping and identification of a candidate gene controlling seed coat color in melon (Cucumis melo L. var. chinensis Pangalo).

KEY MESSAGE: MELO3C019554 encoding a homeobox protein (PHD transcription factor) is a candidate gene that involved in the formation of seed coat color in melon. Seed coat color is related to flavonoid content which is closely related to seed dormancy. According to the genetic analysis of a six-generation population derived from two parents (IC2508 with a yellow seed coat and IC2518 with a brown seed coat), we discovered that the yellow seed coat trait in melon is controlled by a single dominant gene, named CmBS-1. Bulked segregant analysis sequencing (BSA-Seq) revealed that the gene is located at 11,860,000-15,890,000 bp (4.03 Mb) on Chr 6. The F2 population was genotyped using insertion-deletions (InDels), from which cleaved amplified polymorphic sequence (dCAPS) markers were derived to construct a genetic map. The gene was then fine-mapped to a 233.98 kb region containing 12 genes. Based on gene sequence analysis with two parents, we found that the MELO3C019554 gene encoding a homeobox protein (PHD transcription factor) had a nonsynonymous single nucleotide polymorphism (SNP) mutation in the coding sequence (CDS), and the SNP mutation resulted in the conversion of an amino acid (A → T) at residue 534. In addition, MELO3C019554 exhibited lower relative expression levels in the yellow seed coat than in the brown seed coat. Furthermore, we found that MELO3C019554 is related to 12 flavonoid metabolites. Thus, we predicted that MELO3C019554 is a candidate gene controlling seed coat color in melon. The study lays a foundation for further cloning projects and functional analysis of this gene, as well as marker-assisted selection breeding.

QTLs and candidate genes analyses for fruit size under domestication and differentiation in melon (Cucumis melo L.) based on high resolution maps.

BACKGROUND: Melon is a very important horticultural crop produced worldwide with high phenotypic diversity. Fruit size is among the most important domestication and differentiation traits in melon. The molecular mechanisms of fruit size in melon are largely unknown. RESULTS: Two high-density genetic maps were constructed by whole-genome resequencing with two F2 segregating populations (WAP and MAP) derived from two crosses (cultivated agrestis × wild agrestis and cultivated melo × cultivated agrestis). We obtained 1,871,671 and 1,976,589 high quality SNPs that show differences between parents in WAP and MAP. A total of 5138 and 5839 recombination events generated 954 bins in WAP and 1027 bins in MAP with the average size of 321.3 Kb and 301.4 Kb respectively. All bins were mapped onto 12 linkage groups in WAP and MAP. The total lengths of two linkage maps were 904.4 cM (WAP) and 874.5 cM (MAP), covering 86.6% and 87.4% of the melon genome. Two loci for fruit size were identified on chromosome 11 in WAP and chromosome 5 in MAP, respectively. An auxin response factor and a YABBY transcription factor were inferred to be the candidate genes for both loci. CONCLUSION: The high-resolution genetic maps and QTLs analyses for fruit size described here will provide a better understanding the genetic basis of domestication and differentiation, and provide a valuable tool for map-based cloning and molecular marker assisted breeding.

Comprehensive Analysis of Transcriptome and Metabolome Reveals the Flavonoid Metabolic Pathway Is Associated with Fruit Peel Coloration of Melon.

The peel color is an important external quality of melon fruit. To explore the mechanisms of melon peel color formation, we performed an integrated analysis of transcriptome and metabolome with three different fruit peel samples (grey-green 'W', dark-green 'B', and yellow 'H'). A total of 40 differentially expressed flavonoids were identified. Integrated transcriptomic and metabolomic analyses revealed that flavonoid biosynthesis was associated with the fruit peel coloration of melon. Twelve differentially expressed genes regulated flavonoids synthesis. Among them, nine (two 4CL, F3H, three F3'H, IFS, FNS, and FLS) up-regulated genes were involved in the accumulation of flavones, flavanones, flavonols, and isoflavones, and three (2 ANS and UFGT) down-regulated genes were involved in the accumulation of anthocyanins. This study laid a foundation to understand the molecular mechanisms of melon peel coloration by exploring valuable genes and metabolites.

Retention and interference of learned dexterous manipulation: interaction between multiple sensorimotor processes.

An object can be used in multiple contexts, each requiring different hand actions. How the central nervous system builds and maintains memory of such dexterous manipulations remains unclear. We conducted experiments in which human subjects had to learn and recall manipulations performed in two contexts, A and B. Both contexts involved lifting the same L-shaped object whose geometry cued its asymmetrical mass distribution. Correct performance required producing a torque on the vertical handle at object lift onset to prevent it from tilting. The torque direction depended on the context, i.e., object orientation, which was changed by 180° object rotation about a vertical axis. With an A1B1A2 context switching paradigm, subjects learned A1 in the first block of eight trials as indicated by a torque approaching the required one. However, subjects made large errors in anticipating the required torque when switching to B1 immediately after A1 (negative transfer), as well as when they had to recall A1 when switching to A2 after learning B through another block of eight lifts (retrieval interference). Classic sensorimotor learning theories attribute such interferences to multi-rate, multi-state error-driven updates of internal models. However, by systematically changing the interblock break duration and within-block number of trials, our results suggest an alternative explanation underlying interference and retention of dexterous manipulation. Specifically, we identified and quantified through a novel computational model the nonlinear interaction between two sensorimotor mechanisms: a short-lived, context-independent, use-dependent sensorimotor memory and a context-sensitive, error-based learning process.

Coordination between digit forces and positions: interactions between anticipatory and feedback control.

Humans adjust digit forces to compensate for trial-to-trial variability in digit placement during object manipulation, but the underlying control mechanisms remain to be determined. We hypothesized that such digit position/force coordination was achieved by both visually guided feed-forward planning and haptic-based feedback control. The question arises about the time course of the interaction between these two mechanisms. This was tested with a task in which subjects generated torque (± 70 N·mm) on a virtual object to control a cursor moving to target positions to catch a falling ball, using a virtual reality environment and haptic devices. The width of the virtual object was varied between large (L) and small (S). These object widths result in significantly different horizontal digit relative positions and require different digit forces to exert the same task torque. After training, subjects were tested with random sequences of L and S widths with or without visual information about object width. We found that visual cues allowed subjects to plan manipulation forces before contact. In contrast, when visual cues were not available to predict digit positions, subjects implemented a "default" digit force plan that was corrected after digit contact to eventually accomplish the task. The time course of digit forces revealed that force development was delayed in the absence of visual cues. Specifically, the appropriate digit force adjustments were made 250-300 ms after initial object contact. This result supports our hypothesis and further reveals that haptic feedback alone is sufficient to implement digit force-position coordination.

Emerging Medical Technologies and Their Use in Bionic Repair and Human Augmentation.

As both the proportion of older people and the length of life increases globally, a rise in age-related degenerative diseases, disability, and prolonged dependency is projected. However, more sophisticated biomedical materials, as well as an improved understanding of human disease, is forecast to revolutionize the diagnosis and treatment of conditions ranging from osteoarthritis to Alzheimer's disease as well as impact disease prevention. Another, albeit quieter, revolution is also taking place within society: human augmentation. In this context, humans seek to improve themselves, metamorphosing through self-discipline or more recently, through use of emerging medical technologies, with the goal of transcending aging and mortality. In this review, and in the pursuit of improved medical care following aging, disease, disability, or injury, we first highlight cutting-edge and emerging materials-based neuroprosthetic technologies designed to restore limb or organ function. We highlight the potential for these technologies to be utilized to augment human performance beyond the range of natural performance. We discuss and explore the growing social movement of human augmentation and the idea that it is possible and desirable to use emerging technologies to push the boundaries of what it means to be a healthy human into the realm of superhuman performance and intelligence. This potential future capability is contrasted with limitations in the right-to-repair legislation, which may create challenges for patients. Now is the time for continued discussion of the ethical strategies for research, implementation, and long-term device sustainability or repair.

Context-dependent learning interferes with visuomotor transformations for manipulation planning.

How the CNS transforms visual information of object properties into motor commands for manipulation is not well understood. We designed novel apparatus and protocols in which human subjects had to learn manipulations in two different contexts. The first task involved manipulating a U-shaped object that can afford two actions by grasping different parts of the same object. The second task involved manipulating two L-shaped objects that were posed at different orientations. In both experiments, subjects learned the manipulation over consecutive trials in one context before switching to a different context. For both objects and tasks, the visual geometric cues were effective in eliciting anticipatory control with little error at the beginning of learning of the first context. However, subjects failed to use the visual information to the same extent when switching to the second context as sensorimotor memory built through eight consecutive repetitions in the first context exerted a strong interference on subjects' ability to use visual cues again when the context changed. A follow-up experiment where subjects were exposed to a pseudorandom sequence of context switches with the U-shaped object revealed that the interference caused by the preceding context persisted even when subjects switched context after only one trial. Our results suggest that learning generalization of dexterous manipulation is fundamentally limited by context-specific learning of motor actions and competition between vision-based motor planning and sensorimotor memory.

Decoding reach-to-grasp from EEG using classifiers trained with data from the contralateral limb.

Fundamental to human movement is the ability to interact with objects in our environment. How one reaches an object depends on the object's shape and intended interaction afforded by the object, e.g., grasp and transport. Extensive research has revealed that the motor intention of reach-to-grasp can be decoded from cortical activities using EEG signals. The goal of the present study is to determine the extent to which information encoded in the EEG signals is shared between two limbs to enable cross-hand decoding. We performed an experiment in which human subjects (n = 10) were tasked to interact with a novel object with multiple affordances using either right or left hands. The object had two vertical handles attached to a horizontal base. A visual cue instructs what action (lift or touch) and whether the left or right handle should be used for each trial. EEG was recorded and processed from bilateral frontal-central-parietal regions (30 channels). We trained LDA classifiers using data from trials performed by one limb and tested the classification accuracy using data from trials performed by the contralateral limb. We found that the type of hand-object interaction can be decoded with approximately 59 and 69% peak accuracy in the planning and execution stages, respectively. Interestingly, the decoding accuracy of the reaching directions was dependent on how EEG channels in the testing dataset were spatially mirrored, and whether directions were labeled in the extrinsic (object-centered) or intrinsic (body-centered) coordinates.

Learning acquisition of consistent leader-follower relationships depends on implicit haptic interactions.

Are leaders made or born? Leader-follower roles have been well characterized in social science, but they remain somewhat obscure in sensory-motor coordination. Furthermore, it is unknown how and why leader-follower relationships are acquired, including innate versus acquired controversies. We developed a novel asymmetrical coordination task in which two participants (dyad) need to collaborate in transporting a simulated beam while maintaining its horizontal attitude. This experimental paradigm was implemented by twin robotic manipulanda, simulated beam dynamics, haptic interactions, and a projection screen. Clear leader-follower relationships were learned only when strong haptic feedback was introduced. This phenomenon occurred despite participants not being informed that they were interacting with each other and the large number of equally-valid alternative dyadic coordination strategies. We demonstrate the emergence of consistent leader-follower relationships in sensory-motor coordination, and further show that haptic interaction is essential for dyadic co-adaptation. These results provide insights into neural mechanisms responsible for the formation of leader-follower relationships in our society.

Transfer of learned manipulation following changes in degrees of freedom.

The present study was designed to determine whether manipulation learned with a set of digits can be transferred to grips involving a different number of digits, and possible mechanisms underlying such transfer. The goal of the task was to exert a torque and vertical forces on a visually symmetrical object at object lift onset to balance the external torque caused by asymmetrical mass distribution. Subjects learned this manipulation through consecutive practice using one grip type (two or three digits), after which they performed the same task but with another grip type (e.g., after adding or removing one digit, respectively). Subjects were able to switch grip type without compromising the behavioral outcome (i.e., the direction, timing, and magnitude of the torque exerted on the object was unchanged), despite the use of significantly different digit force-position coordination patterns in the two grip types. Our results support the transfer of learning for anticipatory control of manipulation and indicate that the CNS forms an internal model of the manipulation task independent of the effectors that are used to learn it. We propose that sensory information about the new digit placement--resulting from adding or removing a digit immediately after the switch in grip type--plays an important role in the accurate modulation of new digit force distributions. We discuss our results in relation to studies of manipulation reporting lack of learning transfer and propose a theoretical framework that accounts for failure or success of motor learning generalization.

Phloem loading strategies and water relations in trees and herbaceous plants.

Most herbaceous plants employ thermodynamically active mechanisms of phloem loading, whereas in many trees, the mechanism is passive, by diffusion. Considering the different water transport characteristics of herbs and trees, we hypothesized that water relations play a role in the adoption of phloem loading strategies. We measured whole-plant hydraulic conductance (K(p)), osmolality, concentrations of polar metabolites, and key inorganic ions in recently mature leaves of 45 dicotyledonous species at midafternoon. Trees, and the few herbs that load passively, have low K(p), high osmolality, and high concentrations of transport sugars and total polar metabolites. In contrast, herbs that actively load sucrose alone have high K(p), low osmolality, and low concentrations of sugars and total polar metabolites. Solute levels are higher in sugar alcohol-transporting species, both herbs and trees, allowing them to operate at lower leaf water potentials. Polar metabolites are largely responsible for leaf osmolality above a baseline level (approximately 300 mm) contributed by ions. The results suggest that trees must offset low K(p) with high concentrations of foliar transport sugars, providing the motivating force for sugar diffusion and rendering active phloem loading unnecessary. In contrast, the high K(p) of most herbaceous plants allows them to lower sugar concentrations in leaves. This reduces inventory costs and significantly increases growth potential but necessitates active phloem loading. Viewed from this perspective, the elevation of hydraulic conductance marks a major milestone in the evolution of the herbaceous habit, not only by facilitating water transport but also by maximizing carbon use efficiency and growth.

Age-Dependent Upper Limb Myoelectric Control Capability in Typically Developing Children.

Research in EMG-based control of prostheses has mainly utilized adult subjects who have fully developed neuromuscular control. Little is known about children's ability to generate consistent EMG signals necessary to control artificial limbs with multiple degrees of freedom. As a first step to address this gap, experiments were designed to validate and benchmark two experimental protocols that quantify the ability to coordinate forearm muscle contractions in typically developing children. Non-disabled, healthy adults and children participated in our experiments that aimed to measure an individual's ability to use myoelectric control interfaces. In the first experiment, participants performed 8 repetitions of 16 different hand/wrist movements. Using offline classification analysis based on Support Vector Machine, we quantified their ability to consistently produce distinguishable muscle contraction patterns. We demonstrated that children had a smaller number of highly independent movements (can be classified with >90% accuracy) than adults did. The second experiment measured participants' ability to control the position of a cursor on a 1-DoF virtual slide using proportional EMG control with three different visuomotor gain levels. We found that children had higher failure rates and slower average target acquisitions than adults did, primarily due to longer correction times that did not improve over repetitive practice. We also found that the performance in both experiments was age-dependent in children. The results of this study provide novel insights into the technical and empirical basis to better understand neuromuscular development in children with upper-limb loss.

Inter-personal motor interaction is facilitated by hand pairing.

The extent to which hand dominance may influence how each agent contributes to inter-personal coordination remains unknown. In the present study, right-handed human participants performed object balancing tasks either in dyadic conditions with each agent using one hand (left or right), or in bimanual conditions where each agent performed the task individually with both hands. We found that object load was shared between two hands more asymmetrically in dyadic than single-agent conditions. However, hand dominance did not influence how two hands shared the object load. In contrast, hand dominance was a major factor in modulating hand vertical movement speed. Furthermore, the magnitude of internal force produced by two hands against each other correlated with the synchrony between the two hands' movement in dyads. This finding supports the important role of internal force in haptic communication. Importantly, both internal force and movement synchrony were affected by hand dominance of the paired participants. Overall, these results demonstrate, for the first time, that pairing of one dominant and one non-dominant hand may promote asymmetrical roles within a dyad during joint physical interactions. This appears to enable the agent using the dominant hand to actively maintain effective haptic communication and task performance.

Bio-inspired design of a self-aligning, lightweight, and highly-compliant cable-driven knee exoskeleton.

Powered knee exoskeletons have shown potential for mobility restoration and power augmentation. However, the benefits of exoskeletons are partially offset by some design challenges that still limit their positive effects on people. Among them, joint misalignment is a critical aspect mostly because the human knee joint movement is not a fixed-axis rotation. In addition, remarkable mass and stiffness are also limitations. Aiming to minimize joint misalignment, this paper proposes a bio-inspired knee exoskeleton with a joint design that mimics the human knee joint. Moreover, to accomplish a lightweight and high compliance design, a high stiffness cable-tension amplification mechanism is leveraged. Simulation results indicate our design can reduce 49.3 and 71.9% maximum total misalignment for walking and deep squatting activities, respectively. Experiments indicate that the exoskeleton has high compliance (0.4 and 0.1 Nm backdrive torque under unpowered and zero-torque modes, respectively), high control bandwidth (44 Hz), and high control accuracy (1.1 Nm root mean square tracking error, corresponding to 7.3% of the peak torque). This work demonstrates performance improvement compared with state-of-the-art exoskeletons.

Anticipatory planning and control of grasp positions and forces for dexterous two-digit manipulation.

Dexterous object manipulation requires anticipatory control of digit positions and forces. Despite extensive studies on sensorimotor learning of digit forces, how humans learn to coordinate digit positions and forces has never been addressed. Furthermore, the functional role of anticipatory modulation of digit placement to object properties remains to be investigated. We addressed these questions by asking human subjects (12 females, 12 males) to grasp and lift an inverted T-shaped object using precision grip at constrained or self-chosen locations. The task requirement was to minimize object roll during lift. When digit position was not constrained, subjects could have implemented many equally valid digit position-force coordination patterns. However, choice of digit placement might also have resulted in large trial-to-trial variability of digit position, hence challenging the extent to which the CNS could have relied on sensorimotor memories for anticipatory control of digit forces. We hypothesized that subjects would modulate digit placement for optimal force distribution and digit forces as a function of variable digit positions. All subjects learned to minimize object roll within the first three trials, and the unconstrained device was associated with significantly smaller grip forces but larger variability of digit positions. Importantly, however, digit load force modulation compensated for position variability, thus ensuring consistent object roll minimization on each trial. This indicates that subjects learned object manipulation by integrating sensorimotor memories with sensory feedback about digit positions. These results are discussed in the context of motor equivalence and sensorimotor integration of grasp kinematics and kinetics.

Manipulation after object rotation reveals independent sensorimotor memory representations of digit positions and forces.

Planning of object manipulations is dependent on the ability to generate, store, and retrieve sensorimotor memories of previous actions associated with grasped objects. However, the sensorimotor memory representations linking object properties to the planning of grasp are not well understood. Here we use an object rotation task to gain insight into the mechanisms underlying the nature of these sensorimotor memories. We asked subjects to grasp a grip device with an asymmetrical center of mass (CM) anywhere on its vertical surfaces and lift it while minimizing object roll. After subjects learned to minimize object roll by generating a compensatory moment, they were asked to rotate the object 180 degrees about a vertical axis and lift it again. The rotation resulted in changing the direction of external moment opposite to that experienced during the prerotation block. Anticipatory grasp control was quantified by measuring the compensatory moment generated at object lift onset by thumb and index finger forces through their respective application points. On the first postrotation trial, subjects failed to generate a compensatory moment to counter the external moment caused by the new CM location, thus resulting in a large object roll. Nevertheless, after several object rotations subjects reduced object roll on the initial postrotation trials by anticipating the new CM location through the modulation of digit placement but not tangential forces. The differential improvement in modulating these two variables supports the notion of independent memory representations of kinematics and kinetics and is discussed in relation to neural mechanisms underlying visuomotor transformations.

Dimensionality analysis of forearm muscle activation for myoelectric control in transradial amputees.

Establishing a natural communication interface between the user and the terminal device is one of the central challenges of hand neuroprosthetics research. Surface electromyography (EMG) is the most common source of neural signals for interpreting a user's intent in these interfaces. However, how the capacity of EMG generation is affected by various clinical parameters remains largely unknown. In this study, we examined the EMG activity of forearm muscles recorded from 11 transradially amputated subjects who performed a wide range of movements. EMG recordings from 40 able-bodied subjects were also analyzed to provide comparative benchmarks. By using non-negative matrix factorization, we extracted the synergistic EMG patterns for each subject to estimate the dimensionality of muscle control, under the framework of motor synergies. We found that amputees exhibited less than four synergies (with substantial variability related to the length of remaining limb and age), whereas able-bodied subjects commonly demonstrate five or more synergies. The results of this study provide novel insight into the muscle synergy framework and the design of natural myoelectric control interfaces.

Transcriptomic and metabolomic analyses reveal that melatonin promotes melon root development under copper stress by inhibiting jasmonic acid biosynthesis.

Melatonin has been shown to alleviate the effects of abiotic stress and to regulate plant development. Copper, a common heavy metal and soil pollutant, can suppress plant growth and development. In this work, we explored the protective effects of exogenous melatonin on lateral root formation in response to copper stress using melon seeds subjected to three germination treatments: CK1 (control), CK2 (300 µmol/L CuSO4), and MT3 (300 µmol/L melatonin + 300 µmol/L CuSO4). Melatonin pretreatment increased the antioxidant enzyme activities and root vigor, and decreased the proline and malondialdehyde (MDA) contents in the roots of copper-stressed melon seedlings. We then used transcriptomic and metabolomic analyses to explore the mechanisms by which exogenous melatonin protects against copper stress. There were 70 significant differentially expressed genes (DEGs) (28 upregulated, 42 downregulated) and 318 significantly differentially expressed metabolites (DEMs) (168 upregulated, 150 downregulated) between the MT3 and CK2 treatments. Melatonin pretreatment altered the expression of genes related to redox and cell wall formation processes. In addition, we found that members of the AP2/ERF, BBR/BPC, GRAS, and HD-ZIP transcription factor families may have vital roles in lateral root development. Melatonin also increased the level of Glutathione (GSH), which chelates excess Cu2+. The combined transcriptomic and metabolomic analysis revealed DEGs and DEMs involved in jasmonic acid (JA) biosynthesis, including four lipoxygenase-related genes and two metabolites (linoleic acid and lecithin) related to melatonin's alleviation effect on copper toxicity. This research elucidated the molecular mechanisms of melatonin's protective effects in copper-stressed melon.