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Perceived predation risk as a function of predator dietary cues in terrestrial salamanders.

Prey often avoid predator chemical cues, and in aquatic systems, prey may even appraise predation risk via cues associated with the predator's diet. However, this relationship has not been shown for terrestrial predator-prey systems, where the proximity of predators and prey, and the intensity of predator chemical cues in the environment, may be less than in aquatic systems. In the laboratory, we tested behavioural responses (avoidance, habituation and activity) of terrestrial red-backed salamanders, Plethodon cinereus, to chemical cues from garter snakes, Thamnophis sirtalis, fed either red-backed salamanders or earthworms (Lumbricus spp.). We placed salamanders in arenas lined with paper towels pretreated with snake chemicals, and monitored salamander movements during 120 min. Salamanders avoided substrates preconditioned by earthworm-fed (avoidanceX+/-SE=91.1+/-2.5%, N=25) and salamander-fed (95.2+/-2.5%, N=25) snakes, when tested against untreated substrate (control). Salamanders avoided cues from salamander-fed snakes more strongly (75.2+/-5.5%, N=25) than earthworm-fed snakes when subjected to both treatments simultaneously, implying that salamanders were sensitive to predator diet. Salamanders tended to avoid snake substrate more strongly during the last 60 min of a trial, but activity patterns were similar between salamanders exposed exclusively to control substrate versus those subject to snake cues. In another experiment, salamanders failed to avoid cues from dead conspecifics, suggesting that the stronger avoidance of salamander-fed snakes in the previous experiment was not directly due to chemical cues emitted by predator-killed salamanders. Salamanders also did not discriminate between cues from a salamander-fed snake versus a salamander-fed snake that was recently switched (i.e. <14 days) to an earthworm diet. Our results imply that terrestrial salamanders are sensitive to perceived predation risk via by-products of predator diet, and that snake predators rather than dead salamanders may be largely responsible for the release of such chemicals. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Genome size, secondary simplification, and the evolution of the brain in salamanders.

Compared to other vertebrates, even including lampreys and hagfishes in some respects, salamanders exhibit a relatively simple organization of brain and sense organs which is illustrated here using the visual system as an example. The greatest simplicity is found in the bolitoglossine salamanders, yet all bolitoglossines possess highly projectile tongues and rely on vision for survival; furthermore, some species are agile and acrobatic. The unusual features of the visual system of salamanders include small numbers of large neurons, a low degree of morphological differentiation among neurons, a small proportion of myelinated axons in the optic nerve, and an optic tectum consisting essentially of a periventricular cellular layer and a superficial fiber layer. Similar features are found throughout the central nervous system of salamanders and in the lateral line, auditory and olfactory systems as well. Phylogenetic analysis shows that the most parsimonious interpretation of these data is that the simple organization of the brain and sense organs of salamanders was derived secondarily from a more complex ancestral state. We hypothesize that increased genome size has led to simplification of the nervous system in salamanders. Increased genome size appears to have had profound effects on neural development in salamanders, leading to paedomorphosis, the retention of juvenile or even embryonic characteristics into adulthood. In particular, large genome size is associated with large cell size and reduced rates of cell proliferation, migration and differentiation. Secondary simplification has constrained the function of the salamanders' visual system, primarily by increasing cell size and decreasing cell numbers. However, it also has provided an opportunity for the evolution of compensating mechanisms, which have helped to restore or even enhance visual function. Most apparent among the compensatory mechanisms of bolitoglossine salamanders is the presence of well developed ipsilateral retinotectal projections, which apparently enhance depth perception. It is difficult to explain the unusual history of the nervous system in salamanders solely in terms of natural selection and adaptation. Increasing genome size through selfish replication appears to have played a major role in the evolution of salamander brains by imposing functional constraints as well as creating opportunities for overcoming them.

Animals↗

Effects of predator chemical cues and behavioral biorhythms on foraging activity of terrestrial salamanders.

Red-backed salamanders, Plethodon cinereus, show a variety of alarm responses to chemical cues from eastern garter snakes, Thamnophis sirtalis. We measured the foraging activity of red-backed salamanders exposed to water soiled by a garter snake (fed P. cinereus) or to unsoiled water. Salamanders exposed to snake-soiled water showed less foraging activity than salamanders exposed to unsoiled water; therefore, predators could have nonlethal effects on salamander populations. Our results also show additional factors influenced salamander foraging activity. Salamander foraging activity and responsiveness to chemical cues do not appear to have been affected by sex or food deprivation. Salamander foraging activity does appear to have been influenced by activity biorhythms. Foraging activity of animals in both treatments showed a bimodal periodicity that is consistent with natural activity patterns controlled by internal biorhythms. Exposure to snake-soiled water significantly reduced foraging activity during periods of peak foraging activity, but had a subtler effect on foraging activity during natural lulls in activity. We suggest that both activity biorhythms and exposure to chemical cues are important factors affecting salamander foraging behavior.

Animals↗

Paedomorphosis and simplification in the nervous system of salamanders.

Comparative neuroanatomists since Herrick [1914] have been aware of the paradox that the brain of amphibians, especially salamanders, is less complex than one would expect based on their phylogenetic position among the Tetrapoda. Many features of the brain are less differentiated in salamanders than in tetrapod outgroups, including chondrichthyans and bony fishes, and for some brain characters, the salamander brain is even more simple than that of the agnathans. Here, we perform a cladistic analysis on 23 characters of four sensory systems (visual, auditory, lateral line and olfactory) and the brain. Our taxa include myxinoids, lampreys, chondrichthyans, actinopterygians, Latimeria, Neoceratodus and the lepidosirenid lungfishes, amniotes, frogs, caecilians, salamanders and bolitoglossine salamanders. Of the 23 characters we examined, 19 are most parsimoniously interpreted as secondarily simplified in salamanders from a more complex ancestral state, two characters are equally parsimonious under both hypotheses, one character (well developed ipsilateral retinotectal projections) is more complex in bolitoglossine salamanders than in vertebrates generally, and only one character (migration of neurons in the medial pallium) is most parsimoniously interpreted as retention of the plesiomorphically simple condition. Secondary simplification of the salamander brain appears to result from paedomorphosis, or retention of juvenile or embryonic morphology into adulthood. Paedomorphosis is correlated with an increase in genome size, which in turn is positively correlated with cell size, but negatively correlated with cell proliferation and differentiation rates. Available data suggest that, although increasing genome size and paedomorphosis tend to compromise the function of the salamander brain, compensating mechanisms have evolved that may restore or even enhance brain function.

Animals↗

Neurofilaments are prominent in bullfrog olfactory axons but are rarely seen in those of the tiger salamander, Ambystoma tigrinum.

Bullfrog olfactory axons show variable numbers (0-29) of structurally typical neurofilaments (NFs) about 10 nm in diameter. In studies tracking these NFs through serial sections of axons in cross section, they were found to be discontinuous, with a calculated average length of about 118 microns. In contrast to olfactory axons in bullfrogs, those of the tiger salamander Ambystoma trigrinum rarely show NFs. To be certain that the absence of NFs is a specific characteristic of olfactory axons, pieces of salamander spinal cord, optic nerve, and sciatic nerve were examined and found to contain typical NFs. To minimize the possibility that NFs in salamander olfactory axons were degraded or poorly fixed during preparation for electron microscopy, samples were fixed by using a variety of fixative and buffer combinations. To exclude the possibility that proteases degraded NFs during processing, prior to fixation some pieces of olfactory nerve were incubated in physiological saline containing protease inhibitors. Regardless of the preparation method, NFs were generally not seen in salamander olfactory axons. Extracts of salamander olfactory nerve were subjected to SDS-polyacrylamide gel electrophoresis (PAGE) and immunoblotting studies with monoclonal antibodies to the three NF subunit proteins. The immunoblots showed negligible or trace amounts of NF-L (light) and NF-H (heavy), while an NF-M (medium) protein having a molecular mass (Mr) of 160 kD was present in abundance. Extracts of salamander spinal cord, on the other hand, showed all three subunit proteins (with Mrs of 230, 160, and 77 kD). If one assumes that cells assemble structural elements to provide for a given function, the findings suggest that NFs in olfactory neurons of bullfrogs provide a function that may be missing in olfactory neurons of the salamander; the evidence also suggests that the absence of NFs in the salamander may be due to a deficiency in two of the three NF subunit proteins.

Ambystoma↗

Using a GIS model to assess terrestrial salamander response to alternative forest management plans.

A GIS model predicting the spatial distribution of terrestrial salamander abundance based on topography and forest age was developed using parameters derived from the literature. The model was tested by sampling salamander abundance across the full range of site conditions used in the model. A regression of the predictions of our GIS model against these sample data showed that the model has a modest but significant ability to predict both salamander abundance and mass per unit area. The model was used to assess the impacts of alternative management plans for the Hoosier National Forest (Indiana, USA) on salamanders. These plans differed in the spatial delineation of management areas where timber harvest was permitted, and the intensity of timber harvest within those management areas. The spatial pattern of forest openings produced by alternative forest management scenarios based on these plans was projected over 150 years using a timber-harvest simulator (HARVEST). We generated a predictive map of salamander abundance for each scenario over time, and summarized each map by calculating mean salamander abundance and the mean colonization distance (average distance from map cells with low predicted abundance to those with relatively high abundance). Projected salamander abundance was affected more by harvest rate (area harvested each decade) than by the management area boundaries. The alternatives had a varying effect on the mean distance salamanders would have to travel to colonize regenerating stands. Our GIS modeling approach is an example of a spatial analytical tool that could help resource management planners to evaluate the potential ecological impact of management alternatives.

Animals↗

The effects of used motor oil, silt, and the water mold Saprolegnia parasitica on the growth and survival of mole salamanders (genus Ambystoma).

Amphibians appear to be declining worldwide. One cause of their decline may be used crankcase oil which leaks from motor vehicles and washes into ponds. Once in ponds, the oil may either be directly toxic to amphibians, or may indirectly affect them by disrupting food chains. The effects of oil may also be compounded by naturally occurring materials in the water column such as silt. Silt may interfere with respiration across gill surfaces. This study examined the effects of oil and silt on the growth and metamorphosis of larval mole salamanders, Ambystoma opacum and A. tigrinum tigrinum. In Experiment One it examined ponds with and without silty water and oil pollution to determine their suitability as habitats for salamander larvae. In Experiment Two it studied the effects of low levels of oil combined with silt on animals raised in the laboratory and fed prey items not raised in oil. In Experiment Three, it explored the effects of oil at an ecosystem level by raising the salamanders in the field in plastic micromesocosms that mimicked small ponds. Finally, in Experiment Four, in the laboratory, it examined the short-term survival of salamanders in high concentrations of oil. This study found that ponds containing oil and silt produce salamanders of reduced size and weight. Furthermore, while salamanders are relatively robust to the short term effects of large concentrations of used motor oil, oil has deleterious effects on the community and therefore exerts an indirect negative effect on salamanders. In the mi- cro-mesocosms containing oil, salamanders were smaller and weighed less than animals not raised in oil. Furthermore, silt results in reduced growth, earlier metamorphosis, and increased susceptibility to the water mold Saprolegnia parasitica.

Ambystoma↗

Time course of salamander spinal cord regeneration and recovery of swimming: HRP retrograde pathway tracing and kinematic analysis.

The time course of regeneration of supraspinal and descending brachial intraspinal axons was studied using HRP retrograde tracing and kinematic analysis. Five groups of salamanders (10 salamanders/group) received complete thoracic transection 1.0 cm rostral to the hind limbs abolishing swimming. Groups 1-4 recovered for 2, 4, 6, and 8 weeks, respectively, before being filmed to record the animal's ability to swim. After filming, a second transection was made 1.0 cm caudal to the first (at the level of the lumbar enlargement) and HRP was used to label descending axons which had grown past the first lesion. The fifth group was filmed every 2 weeks for 12 weeks before the second transection was made for HRP application. The films were used to perform frame by frame computer analysis of the amplitude and timing of cyclic lateral flexion waves which make up swimming behavior. The earliest return of coordinated swimming behavior was seen 4 weeks after transection (1 of 20 animals). At 6 weeks post-transection, 5 of 10 animals exhibited coordinated swimming. However, the behavior in these animals was subnormal. In the group surviving 8 weeks post-transection, 5 of 10 animals recovered coordinated swimming behavior. In the group that was filmed every 2 weeks, 5 of the 10 salamanders which did recover, exhibited coordinated swimming behavior by the eighth week post-transection. Kinematic analysis of salamanders that exhibited a return of coordinated swimming revealed quantitative differences compared to normal salamanders. While continuous head to tail undulatory waves were present, the propagation time and period were faster than those in normal salamanders. Retransection of the spinal cord abolished coordinated swimming. The numbers and distribution of HRP-labeled supraspinal neurons varied greatly among the animals that displayed recovery of locomotor abilities. In the salamanders examined 6 weeks post-transection the majority of labeled cells were found in medullary nuclei. In recovered salamanders examined 8 and 12 weeks post-transection, HRP-labeled neurons were found in the red nucleus, in the interstitial nucleus of the fasciculus longitudinalis medialis, and in the mesencephalic as well as the medullary reticular neurons. Recovery of coordinated swimming was only observed in salamanders in which descending supraspinal and intraspinal axons were present at the level of the lumbar enlargement (as demonstrated by HRP retrograde labeling). These results indicate that recovery of locomotion is dependent on the reestablishment of descending input and is not a result of changes in spinal reflexes or propagation of electrical activity through the body wall.

Animals↗

Helminth community structure of sympatric eastern American toad, Bufo americanus americanus, northern leopard frog, Rana pipiens, and blue-spotted salamander, Ambystoma laterale, from southeastern Wisconsin.

One-hundred twelve amphibians, including 51 blue-spotted salamanders, Ambystoma laterale, 30 eastern American toads, Bufo americanus americanus, and 31 northern leopard frogs, Rana pipiens, were collected during April-October 1996 from Waukesha County, Wisconsin and examined for helminth parasites. The helminth compound community of this amphibian assemblage consisted of at least 10 species: 9 in American toads, 8 in leopard frogs, and 3 in blue-spotted salamanders. American toads shared 7 species with leopard frogs, and 2 species occurred in all 3 host species. Although there was a high degree of helminth species overlap among these sympatric amphibians, statistically significant differences were found among host species and percent of indirect or direct-life cycle parasites of amphibian species individual component communities (chi2 = 1,015, P < 0.001). American toads had a higher relative abundance of nematodes, 59%, than larval cestodes, 31%, and larval and adult trematodes, 10%, whereas leopard frogs had a higher relative abundance of larval cestodes, 71.3%, and larval and adult trematodes, 25.3%, than nematodes 3.4%. This is related to ecological differences in habitat and dietary preferences between these 2 anuran species. Helminth communities of blue-spotted salamanders were depauperate and were dominated by larval trematodes, 94%, and few nematodes, 6%. Low helminth species richness in this host species is related to this salamander's relatively small host body size, smaller gape size, lower vagility, and more fossorial habitat preference than the other 2 anuran species. Adult leopard frogs and toads had significantly higher mean helminth species richness than metamorphs, but there was no significant difference in mean helminth species richness among adult and metamorph blue-spotted salamanders. Considering adult helminths, the low species richness and low vagility of caudatans as compared with anurans suggest that local factors may be more important in structuring caudatan helminth communities of salamanders than of anuran hosts. Helminth species infecting salamanders may be more clumped in their geographic distribution as compared with anurans, and the role of other hosts and their parasites at the compound community level may be important in structuring helminth communities of salamanders.

Ambystoma↗

Diagnostic and molecular evaluation of three iridovirus-associated salamander mortality events.

In 1998 viruses were isolated from tiger salamander larvae (Ambystoma tigrinum diaboli and A. tigrinum melanostictum) involved in North Dakota and Utah (USA) mortality events and spotted salamander (A. maculatum) larvae in a third event in Maine (USA). Although sympatric caudates and anurans were present at all three sites only ambystomid larvae appeared to be affected. Mortality at the North Dakota site was in the thousands while at the Utah and Maine sites mortality was in the hundreds. Sick larvae were lethargic and slow moving. They swam in circles with obvious buoyancy problems and were unable to remain upright. On the ventral surface, near the gills and hind limbs, red spots or swollen areas were noted. Necropsy findings included: hemorrhages and ulceration of the skin, subcutaneous and intramuscular edema, swollen and pale livers with multifocal hemorrhage, and distended fluid-filled intestines with areas of hemorrhage. Light microscopy revealed intracytoplasmic inclusions, suggestive of a viral infection, in a variety of organs. Electron microscopy of ultra thin sections of the same tissues revealed iridovirus-like particles within the inclusions. These viruses were isolated from a variety of organs, indicating a systemic infection. Representative viral isolates from the three mortality events were characterized using molecular assays. Characterization confirmed that the viral isolates were iridoviruses and that the two tiger salamander isolates were similar and could be distinguished from the spotted salamander isolate. The spotted salamander isolate was similar to frog virus 3, the type species of the genus Ranavirus, while the tiger salamander isolates were not. These data indicate that different species of salamanders can become infected and die in association with different iridoviruses. Challenge assays are required to determine the fish and amphibian host range of these isolates and to assess the susceptibility of tiger and spotted salamanders to heterologous virus isolates.

Amino Acid Sequence↗

Spinal nerves and their bearing on salamander phylogeny.

Examination of the vertebral columns of representatives of all families of salamanders revealed that, in contrast to the condition found in most other vertebrates, salamander spinal nerves of often pass through foramina in the vertebrae. Two kinds of spinal nerve foramina were found: those in the anterior halves of vertebrae, and those in the posterior halves. In addition, many salamanders retain intervertebral nerves. However, within each family or, in a few cases, subfamily there is a characteristic pattern of spinal nerve-vertebral relationships. The first spinal nerve of all salamanders exits through a foramen in the anterior half of the atlas. All more posterior nerves are intervertebral in the families Cryptobranchidae, Hynobiidae and Proteidae. The posterior caudal nerves exit through the posterior halves of the caudal vertebrae in the family Amphiumidae, while in the subfamilies Dicamptodontinae and Rhyacotritoninae all post-sacral nerves exit through the posterior halves of the vertebrae. All but the first three nerves exit through posterior foramina in the family Plethodontidae and the subfamily Ambystomatinae, while all but the first two nerves pass through posterior foramina in the families Salamandridae and Sirenidae. Several fossil salamanders were also examined. These showed that the amphiumid and dicamptodontine-rhyacotritonine nerve patterns had evolved by the Late Cretaceous, and the sirenid pattern had probably evolved by that time. Other Cretaceous genera associated with the Ambystomatoidea still possessed the primitive intervertebral pattern. Using spinal nerve patterns and several other previously described morphological characters, a new hypothesis of the phylogeny of recent and fossil salamanders is presented and compared to earlier proposed phylogenies of the group. A new classification of salamander families is presented.

Animals↗

A connectionist central pattern generator for the aquatic and terrestrial gaits of a simulated salamander.

This article investigates the neural mechanisms underlying salamander locomotion, and develops a biologically plausible connectionist model of a central pattern generator capable of producing the typical aquatic and terrestrial gaits of the salamander. It investigates, in particular, what type of neural circuitry can produce and modulate the two locomotor programs identified within the salamander's spinal cord; namely, a traveling wave of neural activity for swimming and a standing wave for trotting. A two-dimensional biomechanical simulation of the salamander's body is developed whose muscle contraction is determined by the locomotion controller simulated as a leaky-integrator neural network. While the connectivity of the neural circuitry underlying locomotion in the salamander has not been decoded for the moment, this article presents the design of a neural circuit that has a general organization corresponding to that hypothesized by neurobiologists. In particular, the locomotion controller is based on a body central pattern generator (CPG) corresponding to a lamprey-like swimming controller, and is extended with a limb CPG for controlling the salamander's limbs. The complete controller is developed in three stages: first the development of segmental oscillators, second the development of intersegmental coupling for the making of a lamprey-like swimming CPG, and finally the development of the limb CPG and its coupling to the body CPG. A genetic algorithm is used to determine the parameters of the neural circuit for the different stages, given a high-level description of the desired state space trajectories of the different subnetworks. A controller is thus developed that can produce neural activities and locomotion gaits very similar to those observed in the real salamander. By varying the tonic (i.e. non-oscillating) excitation applied to the network, the speed, direction and type of gait can be varied.

Animals↗

Prey detection by vomeronasal chemoreception in a plethodontid salamander.

While chemoreception is involved in a wide variety of salamander behaviors, the chemosensory system that mediates specific behaviors is rarely known. We investigated the role of the vomeronasal system (VNS) in foraging behavior of the red-backed salamander (Plethodon cinereus) by manipulating salamanders' abilities to detect nonvolatile chemical cues emitted by potential prey. Subjects received one of three treatments: (1) impaired vomeronasal system, (2) sham manipulation, and (3) no manipulation. The role of the VNS in mediating foraging on motile prey (Drosophila melanogaster) was investigated under three light conditions (bright, dim, dark). Salamanders with impaired VNSs foraged less efficiently than either of the other experimental groups by displaying the longest latency to attack and the lowest rate of prey capture, especially in the absence of visual cues. A second experiment utilized freshly killed prey to determine whether the VNS takes on added importance in the absence of visual or tactile cues associated with moving prey. Animals with impaired VNSs showed a decreased foraging efficiency on stationary prey under both dark and light conditions. In addition, a mark-recapture study of VNS-impaired and sham salamanders in the field also indicated that salamanders with impaired VNSs consumed fewer stationary prey compared to shams. The study indicates that the VNS plays a substantial role in the foraging behavior of the plethodontid salamander, P. cinereus.

Animals↗

Late Jurassic salamanders from northern China.

With ten extant families, salamanders (urodeles) are one of the three major groups of modern amphibians (lissamphibians). Extant salamanders are often used as a model system to assess fundamental issues of developmental, morphological and biogeographical evolution. Unfortunately, our understanding of these issues has been hampered by the paucity of fossil evidence available to assess the early history of the group. Here we report the discovery of an extraordinary sample of salamander fossils, some with rare soft-tissue impressions, from the Upper Jurassic of China. With over 500 articulated specimens, this assemblage documents the morphological diversity of early urodeles and includes larvae and adults of both neotenic and metamorphosed taxa. Phylogenetic analysis confirms that these salamanders are primitive, and reveals that all basal salamander clades have Asian distributions. This is compelling evidence for an Asian origin of Recent salamanders, as well as for an extensive and early radiation of several major lineages. These discoveries show that the evolution of salamanders has involved phylogenetic and ecological diversification around a body plan that has remained fundamentally stable for over 150 million years.

Animals↗

Environmental conditions prerequisite for complete limb regeneration in the postmetamorphic adult land-phase salamander, Ambystoma.

Historically, postmetamorphic adult land-phase salamanders have been shown to exhibit minimal to nonexistent limb regeneration. Hence, it has been generally accepted that these forms have lost the intrinsic capacity to regenerate a limb. Due to the experimental protocols used, an alternate explanation is also possible: that this intrinsic capacity cannot be expressed when the salamanders are maintained under adverse laboratory environmental conditions. Therefore, this study addresses two questions: 1) What are the optimal environmental conditions for long-term survival of adult land-phase salamanders; and 2) will complete limb regeneration occur in these salamanders if they are maintained under survival conditions. A mixed population of adult Ambystoma were tested under varying conditions of habitat, temperature, humidity, photoperiod, and food source. Complete limb regeneration was possible in 100% of four species of adult postmetamorphic land-phase Ambystoma salamanders given the proper environmental laboratory conditions of a peat moss and potting soil habitat with a controlled temperature of 25 degrees C +/- 5 degrees C, 70% or greater humidity, a 12/12 light/dark photoperiod, a diet including nightcrawlers released into their respective terraria, and an extended observation time of up to 370 days postamputation (dpa). Regeneration was completed during the following range periods for the adult salamanders: A. annulatum, 324 to 370 dpa; A. maculatum, 255 to 300 dpa; A. texanum, 215 to 250 dpa; and A. tigranum, 155 to 180 dpa.

Ambystoma↗

Cytoarchitecture of the tectum mesencephali in salamanders: a Golgi and HRP study.

The tectum mesencephali of salamanders shows a morphology that has long been considered primitive when compared with that of frogs. The alternative hypothesis is that the salamander brain is secondarily simplified. In order to test these two hypotheses, the cytoarchitecture of the tectum and the projections of tectal neurons were studied in 11 species of salamanders. Application of the Golgi method reveals three major morphological types. Type 1 has a very wide dendritic arborization mostly confined to the deep fiber layers, and somata are always located within the most superficial part of the periventricular gray matter. Type 2 possesses a wide to medium-size dendritic arborization. In subtype 2a the somata are located in the uppermost part of the gray, and dendrites always reach the uppermost layer of retinal afferents; in subtype 2b the somata are found in deeper parts of the gray, and dendrites arborize in the deeper layers of retinal afferents; and in subtype 2c the somata are also located in deeper parts, but the wide dendritic arborization is confined to deep fiber layers. Type 3 shows the narrowest dendritic arbors that always reach the upper two tectal fiber layers. The somata are found at any depth of the gray matter. HRP experiments reveal a correlation between morphological differences and the projections of tectal neurons. Type 1- and type 2c-like cells constitute the uncrossed tecto-bulbo-spinal tract, whereas type 1- and type 2a-like cells and migrated large spindle-shaped cells (Salamandra) constitute the crossed tecto-bulbo-spinal tract. Type 3-like neurons project to thalamic, pretectal, and isthmic termination sites. The HRP experiments also demonstrate the existence of two classes of mesencephalic trigeminal cells. A comparison shows that salamanders and frogs possess very similar functional and morphological types of tectal cells. However, tectal cells of salamanders show a "juvenile" morphology, and the number of migrated cells is about 10 times higher in frogs compared to salamanders. Both phenomena are seen as the result of secondary simplification of brain structures in the context of paedomorphosis.

Animals↗