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PubMed1995

Characterization of an ultraviolet photoreception mechanism in the retina of an amphibian, the axolotl (Ambystoma mexicanum).

Spectral sensitivity measurements from the retina of the Mexican salamander Ambystoma mexicanum (the 'axolotl') were used to provide evidence for and to characterize an amphibian ultraviolet-sensitive photoreception mechanism. Electroretinograms (transretinal voltage responses) were recorded from open eye cup preparations to analyze the spectral sensitivity of the retina. Both dark-adapted and white light-adapted preparations exhibited a peak in sensitivity between 360-370 nm. Under selective chromatic adaptation with long-wavelength light the eye cups continued to exhibit a peak sensitivity around 360 nm. These data confirm the presence of ultraviolet-sensitive cones in the retina of Ambystoma salamanders. The possible relevance of these results to the development and behavior of this group of salamanders is briefly discussed.

Ambystoma mexicanum↗
PubMed2005

Testing hypotheses of speciation timing in Dicamptodon copei and Dicamptodon aterrimus (Caudata: Dicamptodontidae).

Giant salamanders of the genus Dicamptodon are members of the mesic forest ecosystem that occurs in the Pacific Northwest of North America. We estimate the phylogeny of the genus to test several hypotheses concerning speciation and the origin of current species distributions. Specifically, we test competing a priori hypotheses of dispersal and vicariance to explain the disjunct inland distribution of the Idaho giant salamander (D. aterrimus) and to test the hypothesis of Pleistocene speciation of Cope's giant salamander (D. copei) using Bayesian hypothesis testing. We determined that available outgroups were too divergent to root the phylogeny effectively, and we calculated Bayesian posterior probabilities for each of the 15 possible root placements for this four-taxon group. This analysis placed the root on the branch leading to D. aterrimus, indicating that current distribution and speciation of D. aterrimus fit the ancient vicariance hypothesis and are attributable to the orogeny of the Cascade Mountains rather than recent inland dispersal. Furthermore, test results indicate that D. copei is distantly related to other coastal lineages and likely originated much earlier than the Pleistocene. These results suggest that speciation within the genus is attributable to ancient geologic events, while more recent Pleistocene glaciation has shaped genetic variation and distributions within the extant species.

Animals↗
PubMed2006

A molecular assessment of phylogenetic relationships and lineage accumulation rates within the family Salamandridae (Amphibia, Caudata).

We examine phylogenetic relationships among salamanders of the family Salamandridae using approximately 2700 bases of new mtDNA sequence data (the tRNALeu, ND1, tRNAIle, tRNAGln, tRNAMet, ND2, tRNATrp, tRNAAla, tRNAAsn, tRNACys, tRNATyr, and COI genes and the origin for light-strand replication) collected from 96 individuals representing 61 of the 66 recognized salamandrid species and outgroups. Phylogenetic analyses using maximum parsimony and Bayesian analysis are performed on the new data alone and combined with previously reported sequences from other parts of the mitochondrial genome. The basal phylogenetic split is a polytomy of lineages ancestral to (1) the Italian newt Salamandrina terdigitata, (2) a strongly supported clade comprising the "true" salamanders (genera Chioglossa, Mertensiella, Lyciasalamandra, and Salamandra), and (3) a strongly supported clade comprising all newts except S. terdigitata. Strongly supported clades within the true salamanders include monophyly of each genus and grouping Chioglossa and Mertensiella as the sister taxon to a clade comprising Lyciasalamandra and Salamandra. Among newts, genera Echinotriton, Pleurodeles, and Tylototriton form a strongly supported clade whose sister taxon comprises the genera Calotriton, Cynops, Euproctus, Neurergus, Notophthalmus, Pachytriton, Paramesotriton, Taricha, and Triturus. Our results strongly support monophyly of all polytypic newt genera except Paramesotriton and Triturus, which appear paraphyletic, and Calotriton, for which only one of the two species is sampled. Other well-supported clades within newts include (1) Asian genera Cynops, Pachytriton, and Paramesotriton, (2) North American genera Notophthalmus and Taricha, (3) the Triturus vulgaris species group, and (4) the Triturus cristatus species group; some additional groupings appear strong in Bayesian but not parsimony analyses. Rates of lineage accumulation through time are evaluated using this nearly comprehensive sampling of salamandrid species-level lineages. Rate of lineage accumulation appears constant throughout salamandrid evolutionary history with no obvious fluctuations associated with origins of morphological or ecological novelties.

Animals↗
PubMed2006

Phylogeographic incongruence of codistributed amphibian species based on small differences in geographic distribution.

Codistributed species may display either congruent phylogeographic patterns, indicating similar responses to a series of shared climatic and geologic events, or discordant patterns, indicating independent responses. This study compares the phylogeographic patterns of two similarly distributed salamander species within the Pacific Northwest of the United States: Cope's giant salamander (Dicamptodon copei) and Van Dyke's salamander (Plethodon vandykei). Previous studies of P. vandykei support two reciprocally monophyletic lineages corresponding to coastal populations, located from the Olympic Mountains to the mouth of the Columbia River, and inland populations within the Cascade Mountains. We hypothesized that D. copei would have a congruent phylogeographic pattern to P. vandykei due to similarity in distribution and dependence upon similar stream and stream-side habitats. We test this hypothesis by estimating the phylogeny of D. copei using approximately 1800bp of mitochondrial DNA and comparing it to that of P. vandykei. Sympatric populations of D. copei and of P. vandykei display an identical phylogeographic pattern, suggesting similar responses within their shared distribution. Populations of D. copei occurring outside the range of P. vandykei displayed high levels of genetic divergence from those sympatric to P. vandykei. Overall, phylogeographic patterns between the two species were ultimately incongruent due to the high divergence of these allopatric populations. These results provide an example of codistributed species displaying overall incongruent phylogeographic patterns while simultaneously displaying congruent patterns within portions of their shared geographic distribution. This pattern demonstrates that a simple dichotomy of congruent and incongruent phylogeographic patterns of codistributed species may be too simplistic and that more complex intermediate patterns can result even from minor differences in species' ranges.

Animals↗
PubMed1977

Fine structural localization of glucose-6-phosphatase activity in the pancreatic islets of two amphibian species (Salamandra salamandra L. and Rana esculenta L.).

The ultrastructural distribution of glucose-6-phosphatase activity has been investigated in the salamander and frog pancreas. In the pancreas of salamander the enzyme was located in the A-cells, while in frog it occurred in all main types of islet cells B-, A-, and D-cells). As a rule, the reaction intensity was higher in the frog islet cells. No reaction was recorded in the exocrine pancreatic tissue of both species. The glucose-6-phosphatase activity was constantly detected in the lumen of rough endoplasmic reticulum and between the nuclear envelopes. Other enzyme localizations, observed especially in the A-cells of the salamander pancreas, were considered possible diffusion artifacts ro remnants of other phosphatase activity. The enzyme distribution in different types of islet cells, as well as its functional significance are discussed in relation to the findings of other authors.

Animals↗
PubMed2001

Systematic status of Hynobius tokyoensis (Amphibia: Urodela) from Aichi Prefecture, Japan: a biochemical survey.

Taxonomic status of a salamander now identified as Hynobius tokyoensis from Aichi Prefecture, Japan, was assessed by allozymic comparisons with its relatives. All three populations of H. tokyoensis from Aichi formed a monophyletic group with populations of H. nebulosus; this group is substantially diverged from a conspecific population from Kanagawa that formed another group with H. lichenatus. In the former group, the population of H. nebulosus from near the type locality was more divergent from the other conspecific populations than were three populations of H. tokyoensis from Aichi. It is quite clear that the salamander from Aichi now identified as H. tokyoensis is genetically much closer to H. nebulosus than to H. tokyoensis from Kanagawa, which is closest geographically to its type locality. This result conforms well to reported results of morphological and genetic studies, and the salamander from Aichi, now called H. tokyoensis, should be identified as H. nebulosus, which is sufficiently diverged from H. tokyoensis to be considered a distinct species.

Algorithms↗
PubMed1983

Use of a monoclonal antibody as a substrate for mature neurons in vitro.

A mouse monoclonal antibody was produced against salamander retinal membranes. It binds to the retina as well as to a wide variety of other salamander tissues and is called Sal-1. Because retinal neurons dissociated from the mature salamander retina adhere poorly to standard substrates, cells were plated onto coverslips previously treated with Sal-1. This previous treatment resulted in a dramatic increase in cell-substrate adhesion. At low concentrations, the antibody had no detectable effect on the light response and fine structure of freshly dissociated rod cells. After several weeks in culture, retinal neurons continued to be attached to Sal-1-treated coverslips and appeared healthy. Furthermore, many cells had extended elaborate cell processes and achieved morphologies characteristic of mature neurons. These results show that antibodies can be used as substrates for the culture of mature neurons. This technique may also prove useful for in vitro studies of a variety of cell systems.

Ambystoma↗
PubMedDate not supplied

Effects of the herbicide atrazine on Ambystoma tigrinum metamorphosis: duration, larval growth, and hormonal response.

We exposed larval tiger salamanders (Ambystoma tigrinum) reared in the laboratory from eggs collected from a prairie wetland in North Dakota to three concentrations of atrazine (0, 75, and 250 micrograms/L) in a static renewal test to determine the pesticide's effect on (1) plasma corticosterone and thyroxine concentrations, (2) larval size, and (3) days-to-stage at stages 2 and 4 of metamorphic climax. We found significant effects of atrazine on each of these response variables. Plasma thyroxine was elevated in both atrazine-exposed groups compared with the control group; plasma corticosterone was depressed in the 75 micrograms/L treatment compared with both the control and 250 micrograms/L treatment. Larvae exposed to 75 micrograms/L atrazine reached stage 4 later but at a size and weight comparable to the control group. By contrast, larvae in the 250 micrograms/L treatment progressed to stage 4 at the same time but at a smaller size and lower weight than larvae in the control group. These results indicate that the herbicide has the potential to influence tiger salamander life history. We present a model consistent with our results, whereby corticosterone and thyroxine interact to regulate metamorphosis of tiger salamanders based on nutrient assimilation and adult fitness.

Ambystoma↗
PubMed2005

Ontogenetic decline of regenerative ability and the stimulation of human regeneration.

Although we cannot regenerate our limbs today, it is likely that when we were embryos we could regenerate many of our tissues, including our limbs. Like other vertebrates, our impressive regenerative abilities were lost during embryogenesis, leaving us with a relatively limited ability to repair tissue damage. In contrast, adult salamanders can reactivate the embryonic regeneration response, and thus they provide the opportunity to discover the principles and mechanisms of tissue and organ regeneration. One important lesson we have learned from salamanders is that regeneration occurs in two steps. While the second step shares the mechanisms of growth control and pattern formation with limb development, the first step is unique and leads to the formation of a regeneration blastema. A second lesson is that connective tissue fibroblasts control regeneration, and that the unique regenerative ability of salamanders (the first step of regeneration) is a consequence of the ability of fibroblasts to dedifferentiate and give rise to blastema cells. Since we all developed limbs as embryos, we all possess the genetic program for making a limb (the second step of regeneration). Therefore, the challenge for inducing limb regeneration in humans is to discover how to induce fibroblast dedifferentiation.

Animals↗
PubMed2000

Mitochondrial evidence on the phylogenetic position of caecilians (Amphibia: Gymnophiona).

The complete nucleotide sequence (17,005 bp) of the mitochondrial genome of the caecilian Typhlonectes natans (Gymnophiona, Amphibia) was determined. This molecule is characterized by two distinctive genomic features: there are seven large 109-bp tandem repeats in the control region, and the sequence for the putative origin of replication of the L strand can potentially fold into two alternative secondary structures (one including part of the tRNA(Cys)). The new sequence data were used to assess the phylogenetic position of caecilians and to gain insights into the origin of living amphibians (frogs, salamanders, and caecilians). Phylogenetic analyses of two data sets-one combining protein-coding genes and the other combining tRNA genes-strongly supported a caecilian + frog clade and, hence, monophyly of modern amphibians. These two data sets could not further resolve relationships among the coelacanth, lungfishes, and tetrapods, but strongly supported diapsid affinities of turtles. Phylogenetic relationships among a larger set of species of frogs, salamanders, and caecilians were estimated with a mitochondrial rRNA data set. Maximum parsimony analysis of this latter data set also recovered monophyly of living amphibians and favored a frog + salamander (Batrachia) relationship. However, bootstrap support was only moderate at these nodes. This is likely due to an extensive among-site rate heterogeneity in the rRNA data set and the narrow window of time in which the three main groups of living amphibians were originated.

Animals↗
PubMed1984

The lipid composition of urodele myelin which lacks hydroxycerebroside and hydroxysulfatide.

The concentrations of cerebrosides and sulfatides were measured in the nervous systems of urodeles and related orders with a high performance liquid chromatographic technique. The peripheral and central nervous systems of all three urodele species, Necturus maculosis (mud puppy, a salamander), Notophthalmus viridescens (eastern red spot newt), and Desmognathus ochropheus (mountain salamander), were found to be completely devoid of alpha-hydroxy fatty acid-containing cerebrosides and sulfatides. All species of reptiles and fish classes close to urodeles contain these galactolipids. The levels of nonhydroxy fatty acid-containing cerebrosides and sulfatides are essentially similar in both urodeles and reptiles. Myelin isolated from Necturus spinal cord had a specific density of 1.07, lighter than mammalian myelin. Except for the absence of hydroxycerebrosides and hydroxysulfatides, the lipid composition of Necturus spinal cord myelin is essentially similar to that of frog and rat myelin. The fatty acids of nonhydroxycerebrosides are rich in monounsaturated homologs of C22-C25, and the sphingoid base consists of both sphinganine and sphingosine. Electron microscopic examination of the sciatic nerve showed that the general structure and interlamellar distances of salamander and newt myelin are identical to those of frog, chameleon, and rat. Necturus myelin, therefore, can be used as a model for the study of the functional and structural role of hydroxygalactolipids.

Animals↗
PubMedDate not supplied

Vertebral development and amphibian evolution.

Amphibians provide an unparalleled opportunity to integrate studies of development and evolution through the investigation of the fossil record of larval stages. The pattern of vertebral development in modern frogs strongly resembles that of Paleozoic labyrinthodonts in the great delay in the ossification of the vertebrae, with the centra forming much later than the neural arches. Slow ossification of the trunk vertebrae in frogs and the absence of ossification in the tail facilitate the rapid loss of the tail during metamorphosis, and may reflect retention of the pattern in their specific Paleozoic ancestors. Salamanders and caecilians ossify their centra at a much earlier stage than frogs, which resembles the condition in Paleozoic lepospondyls. The clearly distinct patterns and rates of vertebral development may indicate phylogenetic separation between the ultimate ancestors of frogs and those of salamanders and caecilians within the early radiation of ancestral tetrapods. This divergence may date from the Lower Carboniferous. Comparison with the molecular regulation of vertebral development described in modern mammals and birds suggests that the rapid chondrification of the centra in salamanders relative to that of frogs may result from the earlier migration of sclerotomal cells expressing Pax1 to the area surrounding the notochord.

Amphibians↗
PubMedDate not supplied

Limb ossification in the Paleozoic branchiosaurid Apateon (Temnospondyli) and the early evolution of preaxial dominance in tetrapod limb development.

Despite the wide range of shapes and sizes that accompany a vast variety of functions, the development of tetrapod limbs follows a conservative pattern of de novo condensation, branching, and segmentation. Development of the zeugopodium and digital arch typically occurs in a posterior to anterior sequence, referred to as postaxial dominance, with a digital sequence of 4-3-5-2-1. The only exception to this pattern in all of living Tetrapoda can be found in salamanders, which display a preaxial dominance in limb development, a de novo condensation of a basale commune (distal carpal/tarsal 1+2) and a precoccial development of digits I and II. These divergent patterns have puzzled researchers for over a century leading to various explanatory hypotheses. Despite many advances in research on tetrapod limb development, the divergent evolution of these two pathways and its causes are still not understood. Based on an extensive ontogenetic series we investigated the pattern of limb development of the 300 Ma old branchiosaurid amphibian Apateon. This revealed a preaxial dominance in limb development that was previously believed to be unique and derived for modern salamanders. The Branchiosauridae are favored as close relatives of extant salamanders in most phylogenetic hypotheses of the highly controversial origins and relationships of extant amphibians. The findings provide new insights into the evolution of developmental pathways in tetrapod limb development, the relationships of modern amphibians with possible Paleozoic antecedents, and their initial timing of divergence.

Animals↗
PubMedDate not supplied

Heterochrony, cannibalism, and the evolution of viviparity in Salamandra salamandra.

The way in which novelties that lead to macroevolutionary events originate is a major question in evolutionary biology, and one that can be addressed using the fire salamander (Salamandra salamandra) as a model system. It is exceptional among amphibians in displaying intraspecific diversity of reproductive strategies. In S. salamandra, two distinct modes of reproduction co-occur: the common mode, ovoviviparity (females giving birth to many small larvae), and a phylogenetically derived reproductive strategy, viviparity (females producing only a few large, fully metamorphosed juveniles, which are nourished maternally). We examine the relationship between heterochronic modifications of the ontogeny and the evolution of the new reproductive mode in the fire salamander. The in vitro development of embryos of ovoviviparous and viviparous salamanders from fertilization to metamorphosis is compared, highlighting the key events that distinguish the two modes of reproduction. We identify the heterochronic events that, together with the intrauterine cannibalistic behavior, characterize the derived viviparous reproductive strategy. The ways in which evolutionary novelties can arise by modification of developmental programs can be studied in S. salamandra. Moreover, the variation in reproductive modes and the associated variation of sequences of development occur in neighboring, conspecific populations. Thus, S. salamandra is a unique biological system in which evolutionary developmental research questions can be addressed at the level of populations.

Animals↗
PubMed1992

The recording of odorant-induced mucosal activity patterns with a voltage-sensitive dye.

1. Fluorescence changes in the dye (WW 781) were monitored at 100 contiguous sites in a 10 x 10-pixel array on the bullfrog and salamander olfactory mucosas every 10 ms in response to odorous stimuli. The odorants were d-limonene, butanol, and amyl acetate, each presented at two concentrations with a 3:1 ratio. 2. The fluorescence signals elicited by these odorous stimuli were nearly identical in shape and time course to the electro-olfactograms (EOGs) recorded from the same animal under identical conditions. Like the EOGs, the fluorescence signals exhibited adaptation and were abolished by both Triton X-100 and ether. There was no measurable fluorescence when the tissue was not stained with the dye, and there was no change in fluorescence when, for stained tissue, nonodorized, humidified air was presented as the stimulus. 3. This technique presumably monitors the same events as the EOG, but has the advantage of simultaneously recording the odorant-induced activity from multiple sites across most of the mucosa. Thus this technique preserves subtle differences heretofore lost by other techniques both in the coarseness of their matrices and in the variability generated by trying to piece together, into one collage, results from numerous presentations given at different times. 4. In all preparations, there was a larger difference in the inherent activity patterns (derived from response magnitudes) between different odorants than between different concentrations of the same odorant. These differences were largest on the mucosa lining the floor of salamander's olfactory sac. d-limonene and butanol gave their largest responses near the internal and external nares, respectively, whereas the responses for amyl acetate were more uniform across the mucosal sheet. In contrast to the salamander, smaller differences were observed for both the roof and the floor of the bullfrog's olfactory sac. For the floor, both amyl acetate and d-limonene elicited similar patterns of response magnitude, whereas butanol differed from each of these odorants by eliciting a larger response on the anteriolateral aspect of the mucosa and a lesser response on the remainder. For the roof, different odorants produced different activity patterns, which had profiles not simply described as regions of maximal and minimal responsiveness. 5. Different inherent activity patterns based on temporal characteristics of the fluorescence responses were also observed for different odorants. Each odorant produced a different pixel-by-pixel pattern for the times at which the responses started and ended. For any given odorant, these temporal patterns paralleled the patterns given by response magnitudes.(ABSTRACT TRUNCATED AT 400 WORDS)

Ambystoma↗
PubMed1998

Neural substrate for motor control of feeding in amphibians.

Descending pathways to premotor/motor centers and their cell groups of origin were studied by means of retrograde biocytin tracing experiments in the frog Discoglossus pictus and the plethodontid salamander Plethodon jordani, which differ remarkably in the structure and function of their feeding apparatus and their feeding strategy. Labeled neurons were found in 30 major cell groups located in the telencephalon, diencephalon, synencephalon, mesencephalon and rhombencephalon. The number and distribution of nuclei are very similar in both species. Furthermore, the descending pathways of these groups of neurons take the same courses inside the medulla oblongata. Axons of most nuclei descend either in the ventromedial or ventrolateral medulla oblongata, and it is concluded that the spatial arrangement of pathways is identical in the species studied. Bilateral electrical stimulation of the optic tectum of the plethodontid salamander Hydromantes italicus elicited strong discharges of short latencies in the hypoglossal nerve. In most hypoglossal motor neurons, excitatory postsynaptic potentials (EPSPs) of short latencies followed paired shocks applied at intervals as short as 3 ms, but showed temporal and spatial facilitation, suggesting that the EPSPs include mono- as well as polysynaptic components. In the ventral white matter, orthodromic single units were found that are candidates for excitatory reticular interneurons. These properties of tectal descending pathways in salamanders strongly differ from those found in toads. Differences in feeding behavior and its control by the premotor/motor networks between the species investigated do not appear to result from anatomically altered input or from a different organization of descending pathways to these premotor/motor centers, but rather from differences in local properties of reticular premotor networks as well as from different effects of neuromodulatory systems.

Animals↗
PubMed1994

Anatomy and forebrain projections of the olfactory and vomeronasal organs in axolotls (Ambystoma mexicanum).

We examined the anatomy of the nasal cavity and forebrain in the axolotl (Ambystoma mexicanum) to determine whether the olfactory and vomeronasal systems are present in this neotenic aquatic salamander. The current study was motivated by two considerations: (a) little is known of the anatomy of the vomeronasal system in aquatic vertebrates, and (b) the presence of both olfactory and vomeronasal systems in larval amphibians has broad implications for the evaluation of these systems in vertebrates. From cresyl-violet-stained sections of snouts we determined that the nasal cavity of axolotls is much like that of terrestrial salamanders. The main chamber of the nasal cavity contains an olfactory epithelium, which is confined to grooves between longitudinal ridges of connective tissue covered in a nonsensory epithelium which lacks goblet cells. Using transmission electron microscopy, we found morphologically distinct olfactory receptor cells: many receptor cells terminate in microvillar dendrites, and fewer terminate in motile cilia with the 9 + 2 microtubule array typical of vertebrate olfactory receptor cells. The ciliated and microvillar cells occur in clusters with little intermingling. Horseradish peroxidase labeling revealed that axons of the olfactory receptor cells terminate in large glomeruli in the main olfactory bulb at the rostral end of the telencephalon. Lateral to the main chamber of the nasal cavity is a diverticulum that is entirely lined with a vomeronasal epithelium containing basal cells, microvillar receptor cells, sustentacular cells that lack specialized processes on the apical surface, and large ciliated cells that may function to move fluid across the vomeronasal epithelium. Unlike the olfactory epithelium, the vomeronasal epithelium lacks Bowman's glands. Using horseradish peroxidase, we determined that the axons of the vomeronasal receptor cells project to the accessory olfactory bulb, a distinct structure dorsal and caudal to the main olfactory bulb. The presence of both olfactory and vomeronasal systems in axolotls and other neotenic salamanders implies that both systems are pleiomorphic in larval amphibians; we therefore suggest that the vomeronasal system may not have originated as an adaptation to terrestrial life.

Ambystoma mexicanum↗
PubMed2005

Muscle fiber angle, segment bulging and architectural gear ratio in segmented musculature.

The anatomical complexity of myomeres and myosepta has made it difficult to develop a comprehensive understanding of the relationship between muscle fiber architecture, connective tissue mechanics, and locomotor function of segmented axial musculature in fishes. The lateral hypaxial musculature (LHM) of salamanders is less anatomically complex and therefore a good system for exploring the basic mechanics of segmented musculature. Here, we derive a mathematical model of the LHM and test our model using sonomicrometry and electromyography during steady swimming in an aquatic salamander, Siren lacertina. The model predicts longitudinal segment strain well, with predicted and measured values differing by less than 5% strain over most of the range. Deviations between predicted and measured results are unbiased and probably result from the salamanders performing slight turns with associated body torsion in our unconstrained trackway swimming experiments. Model simulations of muscle fiber contraction and segment shortening indicate that longitudinal segment strain, for a given amount of muscle fiber strain, increases with increasing initial fiber angle. This increase in architectural gear ratio (AGR = longitudinal strain/fiber strain) is mediated by muscle fiber rotation; the higher the initial fiber angle, the more the fibers rotate during contraction and the higher the AGR. Muscle fiber rotation is additionally impacted by bulging in the dorsoventral (DV) and/or mediolateral (ML) dimensions during longitudinal segment shortening. In segments with obliquely oriented muscle fibers, DV bulging increases muscle fiber rotation, thereby increasing the AGR. Extending the model to include force and work indicates that force decreases with increasing initial muscle fiber angle and increasing DV bulging and that both longitudinal shortening and DV bulging must be included for accurate calculation of segment work.

Animals↗