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At least 1,351 records · Page 75Linked to original sources
PubMed1974

Electrophysiological investigation of GABA-mediated inhibition at the hermit crab neuromuscular junction.

1. The inhibitory neuromuscular junction of the abductor muscle of the large claw of the hermit crab (Eupagurus bernhardus) was investigated using electrophysiological intracellular techniques in order to elucidate further the relative contributions of the pre- and post-synaptic mechanisms of action of GABA and of neural inhibition.2. The electrical constants of the post-synaptic membrane, calculated using the equations for a ;short cable' model, were characteristic of a poorly developed electrical excitability; the specific membrane resistance was usually < 1000 Omega cm(2) and the specific membrane capacitance was > 40 muF/cm(2).3. Stimulation of the excitatory axon to the abductor muscle of the large claw at a frequency of 20 Hz evoked highly facilitating excitatory junction potentials (e.j.p.s); stimulation of the inhibitory axon (60-220 Hz) during the excitatory train elicited inhibition which was manifest as an attenuation of the e.j.p.s.4. The addition of gamma-aminobutyric acid (GABA) to the bathing solution produced a dose-dependent reduction of e.j.p. amplitude and membrane resistance. The inhibitory effect of concentrations (5 x 10(-5) and 1 x 10(-4)M) which caused a 40-75% e.j.p. attenuation could largely be accounted for by a post-synaptic action on membrane conductance.5. Experiments with picrotoxin suggest that presynaptic inhibitory mechanisms have an important role in neurally evoked inhibition.6. Picrotoxin (1-5 x 10(-5)M) effectively blocked neural inhibition and the actions of GABA in this preparation, whereas bicuculline proved to be considerably less potent and therefore less useful as a physiological tool for studying GABA-mediated inhibition in crustacea.

Action Potentials↗
PubMed1975

Crustacean intestinal detergent promotes sterol solubilization.

Although crustacean tissue cholesterol content is high, Crustacea, like other arthropods; are incapable of cholesterol synthesis, and presumably are dependent for maintaining tissue cholesterol stores on the intestinal absorption of ingested sterol. A detergent, N-(N-dodecanoylsarcosyl)taurine, representative of a set of detergents synthesized by the crustacean hepatopancreas and secreted into the intestine, is capable of efficient cholesterol solubilization, and thus of promoting sterol absorption.

Animals↗
PubMed1994

Densities of Vibrio vulnificus in the intestines of fish from the U.S. Gulf Coast.

Densities of Vibrio vulnificus in the intestinal contents of various finfish, oysters, and crabs and in sediment and waters of the U.S. Gulf Coast were determined by the most probable number procedure. Species were identified by enzyme immunoassay. During the winter, densities of V. vulnificus were low, and the organism was isolated more frequently from sheepshead fish than from sediment and seawater. From April to October, V. vulnificus densities were considerably higher (2 to 5 logs) in estuarine fish than in surrounding water, sediment, or nearby oysters and crustacea. Highest densities were found in the intestinal contents of certain bottom-feeding fish (10(8)/100 g), particularly those that consume mollusks and crustaceans. Densities of V. vulnificus in fish that feed primarily on plankton and other finfish were similar to those in oysters, sediment, and crabs (10(5)/100 g). V. vulnificus was found infrequently in offshore fish. The presence of high densities of V. vulnificus in the intestines of common estuarine fish may have both ecological (growth and transport) and public health (food and wound infections) implications.

Alabama↗
PubMed1997

Distribution of Vibrio vulnificus phage in oyster tissues and other estuarine habitats.

Phages lytic to Vibrio vulnificus were found in estuarine waters, sediments, plankton, crustacea, molluscan shellfish, and the intestines of finfish of the U.S. Gulf Coast, but no apparent relationship between densities of V. vulnificus and its phages was observed. Phage diversity and abundance in molluscan shellfish were much greater than in other habitats. V. vulnificus phages isolated from oysters did not lyse other mesophilic bacteria also isolated from oysters. Both V. vulnificus and its phages were found in a variety of oyster tissues and fluids with lowest densities in the hemolymph and mantle fluid. These findings suggest a close ecological relationship between V. vulnificus phages and molluscan shellfish.

Animals↗
PubMed2004

"Candidatus hepatoplasma crinochetorum," a new, stalk-forming lineage of Mollicutes colonizing the midgut glands of a terrestrial isopod.

Uncultivated bacteria that densely colonize the midgut glands (hepatopancreas) of the terrestrial isopod Porcellio scaber (Crustacea: Isopoda) were identified by cloning and sequencing of their 16S rRNA genes. Phylogenetic analysis revealed that these symbionts represent a novel lineage of the Mollicutes and are only distantly related (<82% sequence identity) to members of the Mycoplasmatales and Entomoplasmatales. Fluorescence in situ hybridization with a specific oligonucleotide probe confirmed that the amplified 16S rRNA gene sequences indeed originated from a homogeneous population of symbionts intimately associated with the epithelial surface of the hepatopancreas. The same probe also detected morphotypically identical symbionts in other crinochete isopods. Scanning and transmission electron microscopy revealed uniform spherical bacterial cells without a cell wall, sometimes interacting with the microvilli of the brush border by means of stalk-like cytoplasmic appendages, which also appeared to be involved in cell division through budding. Based on the isolated phylogenetic position and unique cytological properties, the provisional name "Candidatus Hepatoplasma crinochetorum" is proposed for this new taxon of Mollicutes colonizing the hepatopancreas of P. scaber.

Animals↗
PubMed1977

Effect of serum from various animal species on erythrocyte attachment of endotoxins and other bacterial antigens.

Lipopolysaccharide O antigens (endotoxins) and other bacterial antigens readily attach to erythrocytes in vitro. This attachment is prevented by certain mammalian and avian sera. In this study, the inhibitory capacity of sera from lower animals was compared with that of higher animals for a total of 30 species. Antigens and the corresponding antisera included both crude O antigens and purified lipopolysaccharide preparations, the common enterobacterial antigen from Escherichia coli O14, the Vi antigen from Citrobacter ballerup, the polyribose-phosphate antigen from Haemophilus influenzae type b, and the crude teichoic acid antigen from Staphylococcus aureus. Antigen and serum mixtures were incubated at 37 degrees C for 30 min and used for erythrocyte modification; failure of hemagglutination by homologous bacterial antiserum provided evidence of inhibitory capacity. Sera from the classes Mammalia and Aves were very strong inhibitors; those of Reptilia and Osteichthyes were moderate in activity, displaying variation within the classes; those of Amphibia and Chondrichthyes were minimal inhibitors; and those of Merostomata, Crustacea, and Lamellibranchiata displayed questionable or no inhibitory capacity. Inhibitory sera were active with all antigens tested. The findings suggest evolution of inhibitory factors consistent with the theory of two diverging lines of animal phylogeny based on embryological criteria and closely parallel the observations of an endotoxin-altering capacity in vertebrate sera that is not found in invertebrate sera or hemolymph.

Amphibians↗
PubMed1979

Adsorption and growth of Vibrio cholerae on chitin.

Incubation of Vibrio cholerae of O-group serotype 1 with chitin particles resulted in adsorption of vibrios onto chitin; chitin-adsorbed V. cholerae survived exposure to acid better than nonadsorbed vibrios. V. cholerae multiplied in dialyzed chitin suspended in 4.2% NaCl, suggesting that adherence to ingested chitin of crustacea might be of epidemiological significance by providing a substrate for vibrio multiplication as well as protection from gastric acid during stomach transit.

Acids↗
PubMed1992

Ionic currents of morphologically distinct peptidergic neurons in defined culture.

1. The X-organ sinus gland is a major peptidergic neurosecretory system in Crustacea, analogous to the vertebrate hypothalamoneurohypophyseal system. Neuronal somata isolated from the crab (Cardisoma carnifex) X-organ and maintained in primary culture in unconditioned, fully defined medium show immediate regenerative outgrowth. Outgrowth occurring as broad lamellipodia ("veiled") distinguishes neurons consistently showing crustacean hyperglycemic hormone immunoreactivity. Neurons that are immunoreactive against molt-inhibiting hormone and red pigment concentrating hormone antisera give rise to branched neurites ("branched"). 2. The whole-cell variation of the patch-clamp technique was used to study the electrophysiology of these two cell types 24-48 h after plating. Under current clamp, only veiled neurons fired overshooting action potentials either spontaneously or in response to depolarization. 3. Under voltage clamp, net current was predominantly outward. When solutions that suppressed outward current were used, only veiled neurons showed significant inward current. These included a tetrodotoxin (TTX)-sensitive Na current and a slow (time to peak 6-10 ms at 0 mV) Cd-sensitive Ca current (ICa) that was activated at potentials less than -30 mV, was maximal at 0 to +20 mV, and did not reverse at potentials up to +60 mV. 4. In TTX, the form of the Ca current I(V) curve was unchanged by changes of holding potential between -40 and -80 mV, and 75-100% of ICa was available from -40 mV. 5. ICa inactivated slowly and incompletely. Analysis with two-pulse regimes suggested that both inactivation and facilitation mechanisms were present. 6. Outward current was examined in the presence and absence of 0.5 mM Cd2+ (1 microM TTX was always present in the external medium). Cd2+ ions slightly reduced the peak outward current, usually by less than 10% (Vc = -10 to +20 mV; Vh = -80 mV). All additional observations were in the presence of TTX and Cd2+. 7. Both cell types expressed a 4-aminopyridine (4-AP)-sensitive transient current, analogous to IA, and a slower-rising (minimum time to peak 20 ms), sustained current that was partially sensitive to tetraethylammonium, analogous to IK. 8. The mean Vh at which IA was half inactivated was -46 mV, and the mean time constant for removal of inactivation was 46 ms.(ABSTRACT TRUNCATED AT 400 WORDS)

4-Aminopyridine↗
PubMed1994

Presynaptic inhibition is mediated by histamine and GABA in the crustacean escape reaction.

1. Presynaptic inhibition of sensory transmission during the escape reaction in Crustacea has been studied using an in vitro preparation of the crayfish thoracic ganglia. Electrical stimulation of the medial giant fiber mediating the escape reaction induced depolarization in sensory afferent terminals of the coxo-basal chordotonal organ (CBCO). This depolarization was associated with an increase of the membrane conductance and was partially blocked by a gamma-aminobutyric acid (GABA) antagonist, picrotoxin, and by a histamine antagonist, cimetidine. 2. Pressure ejection of histamine on CBCO sensory terminals (CBT) recorded intracellularly, induced a depolarization of the membrane potential accompanied by a large increase of the conductance. Histamine-induced depolarization persisted after blockade of synaptic transmission mediated by Na+ spikes by tetrodotoxin. The amplitude of histamine-induced depolarization increased when negative current was injected into the sensory terminal through the recording electrode. Moreover, injection of chloride into the CBT, which shifts the reversal potential of chloride to a more positive value, resulted in an increase of the amplitude of the histamine-induced depolarization. 3. The existence of separate receptors for GABA and histamine on the CB sensory terminals was demonstrated using two complementary sets of experiments. The first one consisted of using specific blockers of GABA and histamine. Picrotoxin blocked selectively the GABA-induced depolarization of the CB sensory terminals, while it was ineffective in blocking the histamine-induced depolarization. Conversely, cimetidine blocked the histamine-induced depolarization totally, but did not affect the GABA response. The second set of experiments tested for of cross-desensitization between GABA and histamine responses.(ABSTRACT TRUNCATED AT 250 WORDS)

Abdomen↗
PubMed2006

Insights into the immune transcriptome of the shrimp Litopenaeus vannamei: tissue-specific expression profiles and transcriptomic responses to immune challenge.

Infectious disease constitutes a major obstacle to the sustainability of shrimp aquaculture worldwide and a significant threat to natural populations of shrimp and other crustacea. The study of the shrimp immune system, including the response to viral infection, has been hampered by a relative lack of molecular genetic information and of tools suitable for high-throughput assessment of gene expression. In this report, the generation of a cDNA microarray encompassing 2,469 putative unigenes expressed in gills, circulating hemocytes, and hepatopancreas of Litopenaeus vannamei is described. The unigenes printed on the microarray were derived from the analyses of 7,021 expressed sequence tags obtained from standard cDNA libraries as well as from libraries generated by suppression subtractive hybridization, after challenging shrimp with a variety of immune stimuli. The general utility of the cDNA microarray was demonstrated by interrogating the array with labeled RNA from four different shrimp tissues (gills, hemocytes, hepatopancreas, and muscle) and by analyzing the transcriptomic response of shrimp to a lethal challenge with white spot syndrome virus. Our results indicate that white spot syndrome virus infection upregulates (in the hepatopancreas) genes encoding known and potential antimicrobial effectors, while some genes involved in protection from oxidative stress were found to be downregulated by the virus.

Animals↗
PubMed2002

Modulating the modulators: parasites, neuromodulators and host behavioral change.

Neuromodulators can resculpt neural circuits, giving an animal the behavioral flexibility it needs to survive in a complex changing world. This ability, however, provides parasites with a potential mechanism for manipulating host behavior. This paper reviews three invertebrate host-parasite systems to examine whether parasites can change host behavior by secreting neuromodulators. The parasitic wasp, Cotesia congregata, suppresses host feeding partly by inducing the host (Manduca sexta) to increase the octopamine concentration in its hemolymph. The increased octopamine concentration disrupts the motor pattern produced by the frontal ganglion, preventing the ingestion of food. Polymorphus paradoxus (Acanthocephalan) alters the escape behavior of its host, Gammarus lacustris (Crustacea), possibly through an effect on the host's serotonergic system. The trematode Trichobilharzia ocellata inhibits egg-laying in its snail host (Lymnaea stagnalis), partly by inducing the host to secrete schistosomin. Schistosomin decreases electrical excitability of the caudodorsal cells. The parasite also alters gene expression for some neuromodulators within the host's central nervous system. In at least two of these three examples, it appears that the host, not the parasite, produces the neuromodulators that alter host behavior. Producing physiologically potent concentrations of neuromodulators may be energetically expensive for many parasites. Parasites may exploit indirect less energetically expensive methods of altering host behavior. For example, parasites may induce the host's immune system to produce the appropriate neuromodulators. In many parasites, the ability to manipulate host behavior may have evolved from adaptations designed to circumvent the host's immune system. Immune-neural-behavioral connections may be pre-adapted for parasitic manipulation.

Animals↗
PubMed1994

Identification of a cross-reactive allergen (presumably tropomyosin) in shrimp, mite and insects.

A monoclonal antibody to Dermatophagoides pteronyssinus is described that cross-reacts with an IgE-binding antigen present in insects, Crustacea (e.g. shrimp) and other invertebrates. By means of sodium dodecyl sulfate-polyacrylamide gel electrophoresis, gel filtration and immunofluorescence it was shown that this monoclonal antibody presumably recognizes tropomyosin. Tropomyosin was shown to be involved in cross-reactivity between mite, shrimp and insects in shrimp-allergic patients.

Allergens↗
PubMed1993

Wetlands or aquatic ape? Availability of food resources.

A human evolutionary scenario including an ape inhabiting marine wetlands is rational in a number of contexts. The concept is viable ecologically due to the availability of abundant animal foods in a variety of habitats ranging from mangrove forests to coral reefs. The food resources include mollusks, crustacea and fish abundant in wet zones and pools between high and low tide levels. There is seasonal abundance of swarming marsh insects, turtles, eggs and chicks of colonial birds, and occasional beached and dying marine mammals. Some of these foods would provide an enriched source of polyunsaturated essential fatty acids needed for brain development, and thus allow a spiral of increasing brain development, tool utilisation for better food gathering, and vocal communication for group action. The concept is viable also in terms of availability of the ape-human stock in the African Rift Valley, isolated from montane forests during the late Pliocene, and as an adaptive explanation for many of the species-specific human characters not found in other ground living primates.

Animals↗
PubMed2004

cuticleDB: a relational database of Arthropod cuticular proteins.

BACKGROUND: The insect exoskeleton or cuticle is a bi-partite composite of proteins and chitin that provides protective, skeletal and structural functions. Little information is available about the molecular structure of this important complex that exhibits a helicoidal architecture. Scores of sequences of cuticular proteins have been obtained from direct protein sequencing, from cDNAs, and from genomic analyses. Most of these cuticular protein sequences contain motifs found only in arthropod proteins. DESCRIPTION: cuticleDB is a relational database containing all structural proteins of Arthropod cuticle identified to date. Many come from direct sequencing of proteins isolated from cuticle and from sequences from cDNAs that share common features with these authentic cuticular proteins. It also includes proteins from the Drosophila melanogaster and the Anopheles gambiae genomes, that have been predicted to be cuticular proteins, based on a Pfam motif (PF00379) responsible for chitin binding in Arthropod cuticle. The total number of the database entries is 445: 370 derive from insects, 60 from Crustacea and 15 from Chelicerata. The database can be accessed from our web server at http://bioinformatics.biol.uoa.gr/cuticleDB. CONCLUSIONS: CuticleDB was primarily designed to contain correct and full annotation of cuticular protein data. The database will be of help to future genome annotators. Users will be able to test hypotheses for the existence of known and also of yet unknown motifs in cuticular proteins. An analysis of motifs may contribute to understanding how proteins contribute to the physical properties of cuticle as well as to the precise nature of their interaction with chitin.

Amino Acid Motifs↗
PubMed2005

Determination of biological and physicochemical parameters of Artemia franciscana strains in hypersaline environments for aquaculture in the Colombian Caribbean.

BACKGROUND: Artemia (Crustacea, Anostraca), also known as brine shrimp, are typical inhabitants of extreme environments. These hypersaline environments vary considerably in their physicochemical composition, and even their climatic conditions and elevation. Several thalassohaline (marine) environments along the Colombian Caribbean coast were surveyed in order to contribute to the knowledge of brine shrimp biotopes in South America by determining some vital biological and physicochemical parameters for Artemia survival. Additionally, cyst quality tests, biometrical and essential fatty acids analysis were performed to evaluate the economic viability of some of these strains for the aquaculture industry. RESULTS: In addition to the three locations (Galerazamba, Manaure, and Pozos Colorados) reported in the literature three decades ago in the Colombian Caribbean, six new locations were registered (Salina Cero, Kangaru, Tayrona, Bahía Hondita, Warrego and Pusheo). All habitats sampled showed that chloride was the prevailing anion, as expected, because of their thalassohaline origin. There were significant differences in cyst diameter grouping strains in the following manner according to this parameter: 1) San Francisco Bay (SFB-Control, USA), 2) Galerazamba and Tayrona, 3) Kangarú, 4) Manaure, and 5) Salina Cero and Pozos Colorados. Chorion thickness values were smaller in Tayrona, followed by Salina Cero, Galerazamba, Manaure, SFB, Kangarú and Pozos Colorados. There were significant differences in naupliar size, grouping strains as follows (smallest to largest): 1) Galerazamba, 2) Manaure, 3) SFB, Kangarú, and Salina Cero, 4) Pozos Colorados, and 5) Tayrona. Overall, cyst quality analysis conducted on samples from Manaure, Galerazamba, and Salina Cero revealed that all sites exhibited a relatively high number of cysts.g-1. Essential fatty acids (EFA) analysis performed on nauplii from cyst samples from Manaure, Galerazamba, Salina Cero and Tayrona revealed that cysts from all sites exhibited high arachidonic acid:20:4(n-6) (ArA) and eicosapentaenoic acid: 20:5(n-3) (EPA) levels comparable to the control sample (SFB). In contrast, most cysts collected (including SFB) at different locations, and during different months, presented low docosahexaenoic acid: 22:6(n-3) (DHA) levels (Manaure was the only exception with high DHA levels). Some variations in EPA and ArA levels were observed in all sites, contrasting with the much lower DHA levels which remained constant for all locations, except for Manaure which exhibited variable DHA levels. DHA/EPA ratio was overall very low for all sites compared to SFB cysts. All strains had a low DHA/ArA, but a high EPA/ArA ratio, including the control. CONCLUSION: The Colombian A. franciscana habitats analyzed were determined to be thalassohaline, and suitable for A. franciscana development. EFA profiles demonstrated that Tayrona, Galerazamba, Manaure and Salina Cero strains are suitable food for marine fish and crustacean culture because of their high EPA/ArA ratio, but might have to be fortified with DHA rich emulsions depending on the nutritional requirements of the species to be cultured, because of their overall low DHA content. The relatively small nauplii are appropriate for marine larvaeculture. In contrast, the strains from Tayrona, Kangarú, Salina Cero, and Pozos Colorados may be of use but limited to Artemia small biomass production quantities, because of the small surface area of their respective locations; Artemia could be exploited at these locations for local aquaculture applications. In general, cyst quality evaluation for Manaure, Salina Cero and Galerazamba cysts revealed that cysts from these three locations could improve their quality by concentrating efforts on cyst processing techniques. Finally, most locations had great A. franciscana production potential and require different degrees of water quality and/or infrastructure management.

Journal Article↗
PubMed2003

Male-like behavioral patterns and physiological alterations induced by androgenic gland implantation in female crayfish.

The androgenic gland (AG) has been shown to regulate male sexual differentiation and secondary male characteristics in Crustacea. This study presents for the first time in crustaceans evidence for masculinization effects of the AG on reproductive behavior, in addition to morpho-anatomical and physiological effects. AG implantation into immature female red claw crayfish Cherax quadricarinatus inhibited secondary vitellogenesis and development of the ovaries, as well as morphological traits that facilitate maternal egg brooding; it also caused the appearance of secondary male characteristics. However, primary male characteristics and a masculine reproductive system were not developed. In pair encounters, aggression was substantially lower in interactions between AG-implanted and intact females than in interactions within AG-implanted or intact pairs. Moreover, elements of mating behavior, i.e. male courtship displays and false copulations, were exhibited by AG-implanted females in several encounters with intact females. In addition to known morpho-anatomical and physiological effects of the AG in crustaceans, the present study suggests that the AG has novel effects on the neural network that generates social behavior.

Aggression↗
PubMed1998

Retinal location is the key to identifying predators in fiddler crabs (Uca pugilator).

Fiddler crabs (Uca pugilator Bosc; Crustacea, Decapoda) feed and court in open, flat spaces. Their visual system has several putative adaptations to their flat habitat, including the tendency to keep the vertical axis of their eyes perpendicular to the plane of the substratum. It is hypothesized that one of the functions of this behavior is to distinguish accurately predators from conspecifics by aligning the region of the eye sensitive to predators with the region in space in which predators occur. To test this hypothesis, a crab was placed in a glass dish, and a moving stimulus was presented between 20 degrees above and below eye level (horizontal). Stimuli below the crab's horizon hardly ever evoked escape responses, while identical stimuli above the horizon produced escape responses whose frequency varied with the angular size, not the absolute size, of the stimulus. Experiments with artificial horizons showed that it is the position of the stimulus relative to the eye that is important, rather than its position relative to external cues such as the visible horizon, the vertical light gradient or the gravitational horizon. It is concluded that a crab responds to stimuli according to their position relative to its retinal equator. This conclusion is discussed in light of the crabs' natural behavior with respect to supra-horizontal stimuli.

Animals↗
PubMed1978

Intersegmental reflex coordination by a single joint receptor organ (CB) in rock lobster walking legs.

In the decapod Crustacea, Palinurus vulgaris and Fasus lalandii, the reflex influences of one particular proprioceptor organ, the coxo-basal chordotonal organ (CB), on all the muscles operating the proximal and distal joints of the same leg, have been analysed. The distal end of CB was clamped in fine forceps mounted on a servo-controlled stretcher, and CB length changes of 2 mm were applied. Motor unit activity of the different muscles was recorded as electromyograms (EMGs). 1. Two types of proprioceptive reflex evoked by CB length changes have been investigated: (a) resistance reflexes of the two levator and two depressor muscles of the same leg segment, the coxopodite, i.e. 'intrasegmental reflexes', (b) 'intersegmental reflexes' induced in the muscles operating the proximal (T-C) joint of the same leg, and in all eight muscles of the limb segments distat to CB. 2. Both levator muscles respond reflexly to imposed CB stretch (which normally occurs with limb 'depression'), while both depressors respond during CB shortening (or passive "elevation" of the leg). 3. Intersegmentally CB stretch reflexly activates the M-C extensor muscle, and sometimes facilitates the T-C remotor and C-P bender muscles. Shortening of the single CB organ of a leg excites one or two tonic motor units of the T-C promotor and M-C flexor muscles, and also facilitates the remotor, I-M reductor, and the single stretcher-opener excitatory motoneurone. 4. Some of the muscles, particularly the M-C flexor and extensor muscles, are also influenced intersegmentally by the resting length of CB, usually but not invariably in the same direction as for the corresponding dynamic reflexes. The role of the CB chordotonal organ is discussed, with particular consideration of its intersegmental reflex influence on the posture of the entire leg, and on the more complex motor behaviour of locomotion, where it may be specially significant in coordination of the limb in lateral walking. A complex picture of both tonic and dynamic, inra- and intersegmental reflex regulation of the positions and movements of the limb segments, thus emerges.

Animals↗