三疣梭子蟹蜕壳周期肝胰腺、外壳和鳃中钙含量的变化

研究测定了三疣梭子蟹蜕壳周期肝胰腺、鳃和外壳中的钙含量。结果显示,在不同时间,鳃中钙含量变化较小,各时间点差异不显著;蜕壳前期至蜕壳刚完成阶段三疣梭子蟹(软壳蟹)肝胰腺中钙含量最高,蜕壳后第4 d肝胰腺中钙含量最低。三疣梭子蟹外壳中钙含量随着外壳的硬化(钙化)逐渐增加,完全硬化后外壳中钙含量达到最高。研究表明,蜕壳后三疣梭子蟹肝胰腺中钙含量的降低与外壳的钙化之间具有关联性,肝胰腺在三疣梭子蟹蜕壳后钙的储存中起着重要作用,而鳃在钙离子储存中所起的作用可能小于钙转移的作用。     

Construction and nanomechanical properties of the exoskeleton of the barnacle, Amphibalanus reticulatus

Barnacles are some of the major inhabitants of intertidal zones and have calcite-based exoskeleton to anchor and armor their tissues. Structural characterization studies of the specie Ambhibalanus reticulatus were performed to understand the construction of the exoskeleton which forms a light-weight yet stiff structure. The parietal shell is constructed of six compartments to yield a truncated cone geometry, which is neatly fixed onto the basal shell that attaches the organism to the substrate surface. The connections among the different compartments happen through sutured edges and also have chemical interlocking to make the junctions impermeable. Also, the shell parts are furnished with hollow channels reducing the overall mass of the construction. The structure and functions of different parts of the exoskeleton are identified and outlined. Finally, the mechanical properties such as modulus, hardness and fracture toughness of the exoskeleton obtained by indentation techniques are discussed.     

On the morphology and taxonomic position of <Emphasis Type="Italic">Ungulaspis arctoa</Emphasis> (Osteostraci,Agnatha) from the lower devonian of the Severnaya Zemlya archipelago

An additional study of the cephalic shield and trunk scales of the holotype of Ungulaspis arctoa Afanassieva et Karatajūtė-Talimaa has shown that the exoskeleton surface is composed of dentin of at least two generations. Based on new data, the exoskeleton of Ungulaspis is reconstructed. A model for ontogenetic development of the exoskeleton of Ungulaspis arctoa is proposed. The superposition growth of osteostracan exoskeleton is described in detail for the first time using macro-rather than microremains. Using Osteostraci as an example, the existence of two (bipolar and unipolar) types of vertical growth of the exoskeleton in some groups of lower vertebrates is shown. The taxonomic position of Ungulaspis arctoa in the system of Osteostraci is discussed.     

New data on the exoskeleton of the osteostracan genus Aestiaspis (Agnatha) from the Silurian of Saaremaa Island (Estonia) and the Severnaya Zemlya Archipelago (Russia)

The histological structure of exoskeleton of the osteostracan genus Aestiaspis Janvier et Lelievre, 1994 from the Silurian of Saaremaa Island is examined for the first time. The preservation of the material enables the fine exoskeleton structure to be described in detail. The study of Aestiaspis from Saaremaa Island and the Severnaya Zemlya Archipelago has shown structural details of exoskeleton sculpture. The formation of consolidated cephalothoracic shield in the phylogeny of Tremataspidoidei is discussed. The taxonomic position of the genus Aestiaspis in the Osteostraci system is analyzed.     

Gene expression profiling of cuticular proteins across the moult cycle of the crab Portunus pelagicus

Background  

Crustaceans represent an attractive model to study biomineralization and cuticle matrix formation, as these events are precisely timed to occur at certain stages of the moult cycle. Moulting, the process by which crustaceans shed their exoskeleton, involves the partial breakdown of the old exoskeleton and the synthesis of a new cuticle. This cuticle is subdivided into layers, some of which become calcified while others remain uncalcified. The cuticle matrix consists of many different proteins that confer the physical properties, such as pliability, of the exoskeleton.     

The fine structure of a parasitic ciliate Terebrospira during ingestion of the exoskeleton of a shrimp Palaemonetes

The ciliated protozoan, Terebrospira chattoni, invades the exoskeleton of the shrimp, Palaemonetes pugio, eating out long galleries parallel to the surface of the exoskeleton. Solubilization of the exoskeleton occurs around an area of the elaborately infolded surface membrane at the anterior of the organism. Dissolved products of the digestion of the exoskeleton are taken into the body by the formation of coated vesicles at pores in the membrane. The surface membrane that is taken in by pinocytosis is apparently recycled by the introduction into the membrane of organelles implicated in mambrane recycling in other ciliates. Acid phosphatase can be demonstrated on the surface membrane as well as in the endocuticle around the organism.     

A study of strontium,magnesium, and calcium in the environment and exoskeleton of decapod crustaceans,with special reference to Uca burgersi on barbuda,west Indies

Analyses of samples from Barbuda, Leeward Islands, show that the strontium/calcium atom ratio of the exoskeleton of the fiddler crab Uca burgersi Holthuis is proportional to that of the environment, whilst the atom ratio magnesium/calcium of the exoskeleton is fairly constant regardless of the environmental ratio.     

Observations on the Host-Symbiont Relationships of Lagenophrys lunatus Imamura*

An epizoic loricate peritrich, Lagenophrys lunatus Imamura, was found in North Carolina, Virginia, and Maryland on the exoskeleton of the palaemonid shrimp, Palaemonetes pugio, P. intermedius, P. vulgaris, P. paludosus, and Macrobrachium ohione. L. lunatus prefers to attach onto or near the appendages of the host where water movement over the exoskeleton is strongest. The symbionts depend for their food on particles of the host's food and phytoplankton swept to them by these water currents. They attach to other areas of the exoskeleton only when the preferred areas are filled. Trophic individuals are stimulated to undergo a special type of division in advance of the molt of the host that results in the evacuation from the old exoskeleton. Conjugation occurs en masse and has only been observed during molting of the host.     

On the Modes of the Formation of the Exoskeleton in Early Jawless Vertebrates (Osteostraci,Agnatha)

Based on recently obtained original and published data on the fine structure of the external skeleton of osteostracan agnathans (Osteostraci, Agnatha), possible modes of the formation of their hard cover in the course of the horizontal growth of the exoskeleton are characterized. The developmental models for the formation of various configurations of cephalothoracic shields typical for osteostracans are revealed. It is shown that, in the morphogenesis of the hard cover of this group of early vertebrates, a significant part of the variants of the exoskeleton horizontal growth characteristic of early vertebrates are observed.     

Regeneration of the penis in Balanus balanoides (L.)

In Balanus balanoides (L.) the penis is ‘lost’ each autumn. The regeneration of the penis is described. The primordium contains mesenchyme. The pedicel structures develop early as thickened areas of the exoskeleton and give rise to the characteristic structural features of the mature pedicel. Tissue differentiation begins in the pedicel and later extends to the penis proper which becomes annulated. The ductus eventually opens by a terminal invagination of the exoskeleton.     

On the growth and regeneration of the exoskeleton in early jawless vertebrates (Osteostraci,Agnatha)

Based on the study of sculpture of the cephalic shield and histological structure of the exoskeleton in Reticulaspis menneri Afanassieva et Karatajute -Talimaa (Osteostraci, Agnatha) from the Lower Devonian of the Severnaya Zemlya Archipelago, it has been established that, in the ontogeny of the form under study, dentin generation of the exoskeleton may have developed repeatedly, depending on the requirements of the organism. It is established for the first time that, on the shield surface of osteostracans, dentin structures of various types (tubercles, ridges, networks) could be formed both primarily and in subsequent generations with the growth of the exoskeleton. Injury of the integument and, hence, changes in mechanical tension in covering tissues caused activation of the formation of subsequent generations (of dentin) in the areas involved in reparative regeneration of the shield.     

Immunocytochemical localization of actin and tubulin in the integument of land crab (Gecarcinus lateralis) and lobster (Homarus americanus)

The crustacean integument consists of the exoskeleton and underlying epithelium and associated tissues. The epithelium, which is composed of a single layer of cells, is responsible for the cyclical breakdown and synthesis of the exoskeleton associated with molting (ecdysis). During premolt (proecdysis) the epithelial cells lengthen and secrete the two outermost layers (epicuticle and exocuticle) of the new exoskeleton while partially degrading the two innermost layers (endocuticle and membranous layer) of the overlying old exoskeleton. This increased cellular activity is associated with increased protein synthesis and a change in cell shape from cuboidal to columnar. The cytoskeleton, composed of microfilaments (actin) and microtubules (tubulin), plays important roles in the intracellular organization and motility of eukaryotic cells. Immunoblot analysis shows that the land crab exoskeleton contains actin, tubulin, and actin-related proteins (Varadaraj et al. 1996. Gene 171:177-184). In the present study, immunocytochemistry of land crab and lobster integument showed that both proteins were localized in various cell types, including epithelia, connective tissue, tendinal cells, and blood vessels. Muscle immunostained for actin and myosin, but not for tubulin. The membranous layer of land crab (the other layers of the exoskeleton were not examined) and membranous layer and endocuticle of lobster also reacted specifically with anti-beta-actin and anti-alpha-tubulin monoclonal antibodies, but not with an anti-myosin heavy chain antibody. During proecdysis immunolabeling of the membranous layer decreased probably due to protein degradation. The staining intensity for actin and tubulin in the proecdysial epithelium was similar to that in the intermolt (anecdysial) epithelium, suggesting that there was a net accumulation of both proteins proportional to the increase in cellular volume. These results support the previous biochemical analyses and, more specifically, localize actin and tubulin in exoskeletal structures, suggesting that they may serve both intracellular and extracellular functions in crustaceans. J. Exp. Zool. 286:329-342, 2000.     

State Classification and Motion Description for the Lower Extremity Exoskeleton SJTU-EX

A lower extremity exoskeleton, SJTU-EX, is proposed, which mainly aims to help soldiers and workers to support a payload in motions. To solve the issues of the exoskeleton-environment and exoskeleton-human interactions, four types of foot contact are proposed based on their different kinematic characteristics. By the combination of single leg states, sixteen exo- skeleton states are presented using a series of meaningful notations from the topological point of view. The generalized function set （GF set） theory is employed to achieve the kinematic characteristics of the end effectors in different states. Moreover six mathematical formulations of the dynamics of the exoskeleton and its interactions with the human wearer are developed for different exoskeleton states. The applicability and potential of the proposed classification method are demonstrated by analyzing common lower limb motions, which are described concisely by a sequence of notations. Finally, a new concept of characteristic state is put forward to uniquely indicate the type of motion.     

Application of EMG signals for controlling exoskeleton robots.

Exoskeleton robots are mechanical constructions attached to human body parts, containing actuators for influencing human motion. One important application area for exoskeletons is human motion support, for example, for disabled people, including rehabilitation training, and for force enhancement in healthy subjects. This paper surveys two exoskeleton systems developed in our laboratory. The first system is a lower-extremity exoskeleton with one actuated degree of freedom in the knee joint. This system was designed for motion support in disabled people. The second system is an exoskeleton for a human hand with 16 actuated joints, four for each finger. This hand exoskeleton will be used in rehabilitation training after hand surgeries. The application of EMG signals for motion control is presented. An overview of the design and control methods, and first experimental results for the leg exoskeleton are reported.     

Muscle recruitment and coordination with an ankle exoskeleton

Exoskeletons have the potential to assist and augment human performance. Understanding how users adapt their movement and neuromuscular control in response to external assistance is important to inform the design of these devices. The aim of this research was to evaluate changes in muscle recruitment and coordination for ten unimpaired individuals walking with an ankle exoskeleton. We evaluated changes in the activity of individual muscles, cocontraction levels, and synergistic patterns of muscle coordination with increasing exoskeleton work and torque. Participants were able to selectively reduce activity of the ankle plantarflexors with increasing exoskeleton assistance. Increasing exoskeleton net work resulted in greater reductions in muscle activity than increasing exoskeleton torque. Patterns of muscle coordination were not restricted or constrained to synergistic patterns observed during unassisted walking. While three synergies could describe nearly 95% of the variance in electromyography data during unassisted walking, these same synergies could describe only 85–90% of the variance in muscle activity while walking with the exoskeleton. Synergies calculated with the exoskeleton demonstrated greater changes in synergy weights with increasing exoskeleton work versus greater changes in synergy activations with increasing exoskeleton torque. These results support the theory that unimpaired individuals do not exclusively use central pattern generators or other low-level building blocks to coordinate muscle activity, especially when learning a new task or adapting to external assistance, and demonstrate the potential for using exoskeletons to modulate muscle recruitment and coordination patterns for rehabilitation or performance.     

Degradation of barnacle nauplii: implications to chitin regulation in the marine environment

The exoskeleton of most invertebrate larval forms is made of chitin, which is a linear polysaccharide of β (1→4)-linked N-acetylglucosamine (GlcNAc) residues. These larval forms offer extensive body surface for bacterial attachment and colonization. In nature, degradation of chitin involves a cascade of processes brought about by chitinases produced by specific bacteria in the marine environment. Microbial decomposition of larval carcasses serves as an alternate mechanism for nutrient regeneration, elemental cycling and microbial production. The present study was undertaken to assess the influence of chitinase enzyme on the degradation of the nauplii of barnacle, Balanus amphitrite. The survival and abundance of bacteria during the degradation process under different experimental conditions was monitored. To the best of our knowledge, no such study is conducted to understand the degradation of larval exoskeleton using chitinase and its influence on bacteria. An increase in the chitinase activity with increase in temperature was observed. Scanning electron micrographs of chitinase treated nauplii showed scars on the surface of the barnacle nauplii initially and further disruption of the exoskeleton was observed with the increase in the treatment time. Bacterial abundance of the chitinase treated nauplii increased with the increase in enzyme concentration. Pathogenic bacteria such as Vibrio cholerae, V. alginolyticus, V. parahaemolyticus which were initially associated with the exoskeleton were absent after chitinase treatment, however, Bacillus spp. dominated subsequent to chitinase treatment and this might have important implications to marine ecosystem functioning.     

An evolutionary comparative analysis of the medusozoan (Cnidaria) exoskeleton

The benthic polyp phase of Medusozoa (Staurozoa, Cubozoa, Scyphozoa, and Hydrozoa) has endoskeletal or exoskeletal support systems, but their composition, development, and evolution is poorly known. In this contribution the variation in synthesis, structure, and function of the medusozoan exoskeleton was examined. In addition, an evolutionary hypothesis for its origin and diversification is proposed for both extinct and extant medusozoans. We also critically reviewed the literature and included data from our own histological and microstructural analyses of some groups. Chitin is a characteristic component of exoskeleton in Medusozoa, functioning as support, protection, and a reserve for various ions and inorganic and organic molecules, which may persuade biomineralization, resulting in rigid biomineralized exoskeletons. Skeletogenesis in Medusozoa dates back to the Ediacaran, when potentially synergetic biotic, abiotic, and physiological processes resulted in development of rigid structures that became the exoskeleton. Of the many types of exoskeletons that evolved, the corneous (chitin‐protein) exoskeleton predominates today in polyps of medusozoans, with its greatest variation and complexity in the polyps of Hydroidolina. A new type of bilayered exoskeleton in which there is an exosarc complementing the perisarc construction is here described.     

Prey exoskeletons influence the course of gastric evacuation in Atlantic cod Gadus morhua

This study examined the effects of prey exoskeleton characteristics on gastric evacuation patterns in Atlantic cod Gadus morhua. Three distinct stages were highlighted in the gastric evacuation of crustacean prey characterized by a robust exoskeleton. The experiments confirmed that the three shrimp species, Pandalus borealis, Pandalus montagui and Eualus macilentus, and the crab Chionoecetes opilio, were evacuated from the stomach at different rates. The duration of all stages increased with increasing ash (and carbonate) content of the fresh prey. Thickness, chemical composition and morphology of the prey exoskeleton all affected gastric evacuation: duration of initial delay, overall evacuation rate and a decreased evacuation rate at the end of the process. The power exponential function (PEF), with its shape parameter, described the course of evacuation for these prey types well, especially the initial delay. The PEF does not, however, allow describing evacuation by the current stomach content mass independent of meal size, which limits its usefulness in estimating consumption rates of wild G. morhua. To predict and describe gastric evacuation of prey with a robust exoskeleton, it is therefore suggested that the square‐root function be expanded with an initial lag phase, coupled to the mechanistically based cylinder model of gastric evacuation.     

Adaptive Control of Exoskeleton Robots for Periodic Assistive Behaviours Based on EMG Feedback Minimisation

In this paper we propose an exoskeleton control method for adaptive learning of assistive joint torque profiles in periodic tasks. We use human muscle activity as feedback to adapt the assistive joint torque behaviour in a way that the muscle activity is minimised. The user can then relax while the exoskeleton takes over the task execution. If the task is altered and the existing assistive behaviour becomes inadequate, the exoskeleton gradually adapts to the new task execution so that the increased muscle activity caused by the new desired task can be reduced. The advantage of the proposed method is that it does not require biomechanical or dynamical models. Our proposed learning system uses Dynamical Movement Primitives (DMPs) as a trajectory generator and parameters of DMPs are modulated using Locally Weighted Regression. Then, the learning system is combined with adaptive oscillators that determine the phase and frequency of motion according to measured Electromyography (EMG) signals. We tested the method with real robot experiments where subjects wearing an elbow exoskeleton had to move an object of an unknown mass according to a predefined reference motion. We further evaluated the proposed approach on a whole-arm exoskeleton to show that it is able to adaptively derive assistive torques even for multiple-joint motion.     

Effects of a passive exoskeleton on the mechanical loading of the low back in static holding tasks

With mechanical loading as the main risk factor for LBP in mind, exoskeletons are designed to reduce the load on the back by taking over a part of the required moment. The present study assessed the effect of a passive exoskeleton on back and abdominal muscle activation, hip and lumbar flexion and on the contribution of both the human and the exoskeleton to the L5/S1 net moment, during static bending at five different hand heights. Two configurations of the exoskeleton (LOW & HIGH) differing in angle-torque characteristics were tested. L5/S1 moments generated by the subjects were significantly reduced (15–20% for the most effective type) at all hand heights. LOW generated 4–11 Nm more support than HIGH at 50%, 25% and 0% upright stance hand height and HIGH generated 4–5 Nm more support than LOW at 100% and 75%. Significant reductions (11–57%) in back muscle activity were found compared to WITHOUT for both exoskeletons for some conditions. However, EMG reductions compared to WITHOUT were highly variable across subjects and not always significant. The device allowed for substantial lumbar bending (up to 70°) so that a number of participants showed the flexion-relaxation phenomenon, which prevented further reduction of back EMG by the device and even an increase from 2% to 6% MVC in abdominal activity at 25% hand height. These results indicate that flexion relaxation and its interindividual variation should be considered in future exoskeleton developments.