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PubMed1964

THE NATURE OF WATER TRANSPORT ACROSS FROG SKIN.

A method has been developed for determining simultaneously shortcircuit currents and net water fluxes across frog skin. The basis of the water flux measurement is the determination of changes in weight of a plastic chamber containing the skin and external solution. The accuracy of this method permits net water flows larger than 0.5 mg cm(-2)hr.(-1) to be detected, and the apparatus has been used to investigate the relationship between active Na transport and non-osmotic water flow across the skin. Measurement of Na transport and net water influx across completely short-circuited skins provides no good correlation between the two flows. However, skins exhibiting no net water movement in sulfate Ringer displayed an apparent electroosmotic flow of about 40 water molecules per Na ion when depolarizing current densities of 50 and 100 muA cm(-2) are used. It is concluded from this and other evidence that the net water influx across frog skin may be partially electroosmotic in character and that there remains another component of water flow unrelated to active Na transport. A theoretical model, based on irreversible thermodynamics, has been developed to explain the non-osmotic water flow across frog skin.

Animals↗
PubMed1963

SODIUM FLUXES IN DESHEATHED FROG SCIATIC NERVE.

Desheathed frog (R. pipiens) sciatic nerves were soaked in solutions that contained ouabain or NaN(3), in K-free solutions, or in K- and Ca-free solutions. Then, the nerves were allowed to recover in standard Ringer's solution. At various times during the soaking periods, some nerves were analyzed for Na and K, and estimates were made of the influx and efflux of Na(22). While a nerve was soaking in any one of the experimental solutions, the Na(22) influx was increased, the Na content was rising, and the K content was falling by an equivalent amount. The rate coefficient for Na(22) efflux was reduced by about 25 per cent by 0.05 mM ouabain and by about 50 per cent by 5 mM NaN(3). Potassium-free solutions had little effect on the rate coefficient. It was concluded that the efflux of Na from frog nerve is dependent on the metabolism, but not on the external concentration of K. The equimolar exchange that is characteristic of the net movements of Na and K in frog nerve may not be due to the presence of a tightly coupled Na-K exchange pump, but may represent a constraint imposed by the requirement for electroneutrality.

Animals↗
PubMed1964

ELECTRICAL EXCITABILITY OF ISOLATED FROG SKIN AND TOAD BLADDER.

When current of proper polarity and sufficient intensity is passed across isolated frog skin or toad bladder, an action potential of about 200 mv and 10 msec. duration with a sharp threshold and refractory period of several seconds' duration is elicited. Interruption of current during the action potential abolishes the response, and, as shown by appropriate bridge measurements, this occurs because the action potential results from resistance variations during the current flow. The ionic composition of the medium bathing the frog skin was varied, and it was found that the response is relatively insensitive to changes in the solution bathing the inner surface, but rapidly and reversibly affected by changes in the outer solution, particularly by replacement of sodium with potassium and by variations of calcium concentration. It was also observed that the resistance of the skin and action potential across it are reversibly altered by metabolic inhibitors and that these alterations occur independently of any changes in the intrinsic EMF of the system. From the finding that the action potential across frog skin and toad bladder results from a time-variant resistance, it is argued that this same phenomenon can be the basis of electrical excitability in general. This would attribute physical significance to the equivalent circuit commonly employed to represent the plasma membrane; i.e., the plasma membrane would be a mosaic structure of spatially separate permselective regions.

Action Potentials↗
PubMed1996

Developmental expression of a neuron-specific beta-tubulin in frog (Xenopus laevis): a marker for growing axons during the embryonic period.

In mammals, there are seven classes of beta-tubulin genes, one of which, class III, is neuron specific. Using class-specific monoclonal antibodies, class III beta-tubulin protein could not be detected in frog embryos or in adults with either Western blotting or immunohistochemical techniques. In contrast, the class II beta-tubulin protein, which is predominant in mammalian brain but is also expressed in other tissues, is expressed only in neurons in frog embryos. Protein was detected only in neurons from late stages of neural tube closure through premetamorphic stages. At stages 21-28, the pioneering axons of Rohon-Beard, commissural, primary motor, and trigeminal ganglion neurons were distinctly stained in the axon scaffolds that they formed in the embryonic brain and the peripheral mesenchyme. Nonneuronal cells, both outside the nervous system and within it (e.g., radial glia, Müller glia, roof plate, and floor plate cells) were not immunoreactive. Throughout swimming and premetamorphic stages, neuronal cells in all brain regions became immunoreactive as they differentiated and extended axons. Whereas many embryonic neurons became postmitotic during gastrulation stages, neurons expressed detectable levels of class II beta-tubulin protein only beginning at the onset of overt axon outgrowth. These observations demonstrate that the neuron-specific beta-tubulin in frog is a different gene from that in mammals, and its protein product is detectable at the time of axonogenesis rather than neurogenesis.

Amino Acid Sequence↗
PubMed1999

Characterization and localization of pituitary adenylate cyclase-activating polypeptide (PACAP) binding sites in the brain of the frog Rana ridibunda.

The biochemical characteristics and the distribution of pituitary adenylate cyclase-activating polypeptide (PACAP) binding sites have been investigated in the brain of the frog Rana ridibunda by using [(125)I]PACAP27 as a radioligand. Membrane-binding studies revealed the existence of high-affinity receptors for frog PACAP38 and PACAP27. In contrast, the [Des-His(1)]PACAP38 analogue had a much lower affinity and vasoactive intestinal polypeptide did not produce any displacement of the binding. Autoradiographic labeling of frozen brain sections revealed that the highest concentrations of PACAP receptors were located in the olfactory bulb, pallium, striatum, habenular nuclei, ventromedial thalamic nucleus, corpus geniculatum, posterior tubercle, dorsal part of the magnocellular preoptic nucleus, tectum, and the molecular cell layer of the cerebellum. Moderate binding was observed in the septum, in most parts of the thalamus, the dorsal hypothalamic nucleus, the median eminence, the ventral nuclei of the tegmentum, the torus semicircularis, and the interpeduncular and isthmi nuclei. The present data provide the first biochemical characterization and anatomic distribution of PACAP binding sites in the brain of a nonmammalian vertebrate species. The widespread distribution of specific PACAP receptors in the frog brain suggests that the peptide does not act solely as a hypophysiotropic factor, but likely fulfills neurotransmitter functions, neuromodulator functions, or both.

Animals↗
PubMed2000

Structure and distribution of the mRNAs encoding pituitary adenylate cyclase-activating polypeptide and growth hormone-releasing hormone-like peptide in the frog, Rana ridibunda.

The structure of the neuropeptide pituitary adenylate cyclase-activating polypeptide (PACAP) has been characterized in several species including protochordates, fish, amphibians, birds, and mammals. Although PACAP has been shown to stimulate frog pituitary and adrenal cell activity, the structure of the PACAP precursor and the expression of its gene have not yet been reported in any amphibian species. In this study, we have characterized two cDNA variants encoding PACAP of the frog Rana ridibunda, one of which encodes a second peptide exhibiting strong homologies to growth hormone-releasing hormone (GHRH) of fish and mammals. Northern blot and reverse transcriptase-polymerase chain reaction (RT-PCR) analyses revealed that PACAP/GHRH-like peptide mRNAs are predominantly expressed in the brain and spinal cord and, to a lesser extent, in the neurointermediate lobe of the pituitary. Other tissues including the testis and the distal lobe of the pituitary do not express the PACAP precursor gene. The distribution of PACAP/GHRH-like peptide mRNAs in the frog brain has been determined by in situ hybridization histochemistry. High levels of expression were found in the accessory olfactory bulb, the distal pallium, the ventral part of the magnocellular preoptic nucleus, the ventral hypothalamic nucleus, the posterior tuberculum, and the ventral habenular nucleus. These data contribute to the understanding of the evolution of the PACAP and GHRH genes in vertebrates and provide the anatomical bases to elucidate the roles of PACAP and the GHRH-like peptide in amphibians.

Amino Acid Sequence↗
PubMed1997

Hormonal induction of thumb pads and the evolution of secondary sexual characteristics of the Southeast Asian fanged frog, Rana blythii.

The fanged frogs of Southeast Asia do not express most of the hormone-dependent secondary sexual characteristics such as thumb pads that are common to other ranid frogs. At the same point in the evolutionary history of the group that these androgen-mediated characteristics are lost, male parental care first evolves. This behavior is often correlated with low androgen levels. Prior work indicates that in one of the fanged frogs, Rana blythii, adult males have low androgen levels compared to North Temperate species of Rana. This leads to the question of whether these low androgen levels are related to the unusual male parental care and the lack of expression of the thumb pad and other hormone-dependent secondary sexual characteristics in this species. We tested that hypothesis by examining the effects of exogenous dihydrotestosterone supplements on the expression of thumb pads in Rana blythii. Dihydrotestosterone injections appear to stimulate the expression of the thumb pad in R. blythii. These results support the hypothesis that low androgen levels are involved in the loss of the thumb pad in R. blythii. This work provides an example of how mapping characters on phylogenies can be a powerful approach for gaining insights into proximate physiological mechanisms of selection at the evolutionary level.

Animals↗
PubMed1999

Sex-determining mechanism in Buergeria buergeri (Anura, Rhacophoridae). III. Does the ZZW triploid frog become female or male?

Both triploids and gynogenetic diploids (GDs) were produced to clarify the relationship between the sex-chromosome constitution and the expression of sex in the common bell-ring frog, Buergeria buergeri. The sex differentiation of triploids in B. buergeri is quite remarkable. Triploid frogs consisted of three sex genotypes, ZZZ, ZWW and ZZW. All ZZZ triploids were males, and all ZWW triploids were females. It is very interesting that half of the ZZW triploids became female, and the other half became male. The GD frogs consisted of two sex genotypes, ZW and ZZ, which did not differ from the controls in sex differentiation. Since the ratios of ZZ and ZW eggs were significantly different among female parents, it is assumed that most (approximately 80-90%) of the eggs made pre-reductional division in some females and post-reductional division in others during meiosis. It seems that ZW eggs were produced by the occurrence of recombination between the centromere and the sex-determining genes in B. buergeri. It was also found that the number of Z chromosomes in each cell of these triploids and GDs agreed with that of the nucleoli in each cell.

Animals↗
PubMed1996

Ultra-high-resolution scanning electron microscopy of the continuity of cytoplasmic and luminal membranes in frog oxyntic cells.

BACKGROUND: Despite numerous previous studies, the presumed continuity of the luminal and tubulovesicular membranes in frog oxyntic (oxyntico-peptic) cells remains to be convincingly demonstrated. This study was undertaken to clarify this question by ultra-high-resolution scanning electron microscopy of specially prepared frog stomach specimens before and during histamine stimulation. METHODS: Fasted Japanese meadow frogs stimulated with histamine were used. Oxyntic cell cytoplasmic matrix was removed by the aldehyde-osmium-DMSO-osmium maceration procedure and impregnated with osmium-hydrazine. Specimens were examined without metal coating. RESULTS: In the resting oxyntic cell, the luminal membrane had closely packed surface folds forming a rather flat surface with a few short microvilli. In the cytoplasm, flattened 200-500 nm vesicles were interconnected by numerous slender 20-60 nm tubules forming the tubulovesicular network. Occasional slender tubular branches were found in continuity with the luminal membrane. After histamine stimulation, the number and length of microvilli and surface folds increased, whereas the tubulovesicular membrane system decreased. Sites of clear continuity between the luminal and tubulovesicular membranes were not abundant but were clearly demonstrated in histamine-stimulated oxyntic cells. The small size of the tubules connecting the tubulovesicular system to the plasma membrane renders this observation by transmission electron microscopy very difficult. In these specimens, the clear continuity of the tubulovesicular network to the luminal plasma membrane became more evident. CONCLUSIONS: This study confirms previous findings of increased luminal membrane and depletion of the tubulovesicle system. The demonstration of continuity between these two compartments in our SEM preparations supports the hypothesis of direct transfer of tubulovesicular membrane to oxyntic cell luminal secretory membrane.

Animals↗
PubMed1996

Hopping and swimming in the leopard frog, Rana pipiens: II. A comparison of muscle activities.

Electromyography (EMG) was used to examine muscle activity of the major hip, knee, and ankle extensors during both hopping and swimming in leopard frogs. Chronic EMG electrodes were implanted for periods of 7-10 days. This permitted us to record EMG activities during both hopping and swimming from the same electrode, allowing a direct comparison of the timing and amplitudes of muscle activity between the two behaviors. We could then relate these activities to the kinematics of locomotion. In both behaviors, all three extensors were synchronously activated 30-50 ms before limb extension began. However, the hip extensor turned on relatively earlier in hopping than in swimming when on time was expressed as percent of stride. The hip and knee extensors were activated relatively longer in hopping and the ankle extensor relatively longer in swimming. The amplitudes of the rectified, integrated EMG signals were roughly twice as large in hopping as in swimming for all three muscles, supporting the notion that propulsion in hopping requires more force than in swimming. The EMG burst durations differed little between the muscles or, in relative duration, between the behaviors. As has been found in other quadrupeds, the EMG bursts began before visible movement and ceased at or before hindlimb extension was completed. In our animals, however, we found a consistent, low level (10-30% of maximum amplitude) of EMG activity that continued 60-200 ms past the end of the burst and into the suspension periods in both hopping and swimming. We hypothesize that this unusual activity may be present in frogs so that the hind limb remains aero(hydro)dynamically stable as the frog arches through its leap or glides in swimming following completed limb extension. Thus, the timing and pattern of the EMG bursts are consistent with those present in other tetrapods and support conservatism of neural control. However, the prolonged low-level activity suggests flexibility in the control pattern and variation according to specific behaviors.

Animals↗
PubMed1996

Hopping and swimming in the leopard frog, Rana pipiens: I. Step cycles and kinematics.

This study presents a model for the step cycle patterns used during both hopping and swimming by the leopard frog, Rana pipiens. The two behaviors are essentially similar in movement pattern and in the ways they are modified from quadrupedal gaits. In hopping, there is marked hind limb extension throughout stance. The swing begins with a suspension equivalent to the leap that occurs in a galloping or bounding quadruped. Following suspension, as the frog descends from the apex of its leap, the hind limbs remain posterior and in line with the spine while they flex. Near the end of flexion, there is a rapid downward rotation of the hindquarters to bring the hind feet underneath the body. This movement utilizes the planted forelimb as a pivot. A similar pattern of movement occurs in swimming; the stance (propulsion) phase involves extension at all hind limb joints. The swing (recovery) phase begins with the hind feet fully extended and includes a protracted gliding phase, equivalent to the suspension in the hop. The hind limb then recovers to its initial position during a flexion phase. Since there is no landing and the hind limbs remain lateral rather than ventral to the pelvis, less flexion occurs in the spine or the limb joints. In both behaviors, the extensor muscles of hip (M. semimembranosus), knee (M. cruralis), and ankle (M. plantaris longus) achieve their longest lengths, when they likely can produce near maximal force, at the beginning of extension. All three muscles shorten during extension, but, because they are multiple-joint muscles, the amount of shortening is relatively small (approximately 15%). Hopping and swimming in frogs are comparable asymmetrical gaits with the same relative contact intervals (25% of stride). The step cycles in both gaits are modified from quadrupedal locomotion in the same ways: by 1) loss of knee and ankle extension toward the ground prior to landing (or end of flexion in swimming), 2) loss of a yield phase on landing (or end of flexion in swimming), and 3) inclusion of extended suspensions in both gaits.

Animals↗
PubMed1999

Description of the chondrocranium and osteogenesis of the Chacoan burrowing frog, Chacophrys pierotti (Anura: Leptodactylidae).

The larval chondrocranium of the large-headed leptodactylid frog, Chacophrys pierotti (Ceratophryinae), is described in detail. Descriptions include the ontogeny of the chondrocranium and osteogenesis of the cranial skeleton. The chondrocranium of C. pierotti is profoundly different from the chondrocrania previously described for the other genera of the Ceratophryinae (Ceratophrys and Lepidobatrachus). The chondrocranium of Chacophrys is longer than wide and not particularly robust or laterally expanded; that of Ceratophrys is very robust, whereas the chondrocranium of Lepidobatrachus is widely expanded laterally. These differences are particularly apparent in the elements associated with the jaw (i.e., suprarostral, infrarostral, Meckel's cartilage, palatoquadrate, cornua trabeculae), which are robust in Ceratophrys and thin and elongate in Lepidobatrachus. Unlike Ceratophrys and Lepidobatrachus, which possess highly specialized carnivorous larva, the chondrocranium of Chacophrys more closely resembles the typical microphagous herbivore morphology characteristic of other leptodactylid frogs for which the chondrocrania are known. These data suggest that Chacophrys is the basal taxon within the monophyletic Ceratophryinae. The ontogeny of the chondrocranium of Chacophrys, as well as the cranial ossification sequence, do not differ greatly from those described for Ceratophrys. Detailed descriptions of the ontogeny of the chondrocranium and the bony skeleton are needed for additional taxa within the Ceratophryinae (especially Lepidobatrachus). Such descriptive ontogenetic studies promise new insight into the phylogeny and morphological evolution of this remarkable group of large-headed frogs.

Animals↗
PubMed2001

Immunohistochemical localization and biochemical characterization of hypocretin/orexin-related peptides in the central nervous system of the frog Rana ridibunda.

In the present study, we have investigated the distribution and biochemical characteristics of hypocretin (hcrt) -like immunoreactivity in the central nervous system (CNS) of the frog Rana ridibunda by using an antiserum directed against rat hcrt2. Immunoreactive cell bodies were only detected in four diencephalic nuclei, including the anterior preoptic area and the suprachiasmatic, magnocellular, and ventral hypothalamic nuclei. In contrast, hcrt2-immunoreactive fibers were widely distributed throughout the frog CNS. In particular, a high density of hcrt-positive fibers was detected in several areas of the telencephalon, including the olfactory bulb, the nucleus of the diagonal band of Broca, and the amygdala. A dense network of hcrt-containing fibers was observed in all thalamic and hypothalamic nuclei. A low to moderate density of immunoreactive fibers was also found in the mesencephalon, rhombencephalon, and spinal cord. Reversed-phase high performance liquid chromatography analysis of frog brain extracts revealed that hcrt2-immunoreactive material eluted as two peaks, the major one exhibiting the same retention time as synthetic rat hcrt2. The present data provide the first detailed mapping of the hcrt neuronal system in the CNS of a nonmammalian vertebrate. The occurrence of hcrt-containing cell bodies in the hypothalamus and the widespread distribution of hcrt-immunoreactive fibers throughout the brain and spinal cord suggest that, in amphibians, hcrts may exert neuroendocrine, neurotransmitter, and/or neuromodulator activities.

Animals↗
PubMed2000

The antibiotic and anticancer active aurein peptides from the Australian Bell Frogs Litoria aurea and Litoria raniformis. Part 2. Sequence determination using electrospray mass spectrometry.

Sixteen aurein peptides are present in the host defence secretion from the granular dorsal glands of the Green and Golden Bell Frog Litoria aureus and seventeen from those of the related Southern Bell Frog Litoria raniformis. All peptides have been sequenced using a combination of electrospray mass spectrometry and Lys-C digestion, with each sequence confirmed by automated Edman sequencing. The peptides are named in five groups, viz. aureins 1-5. Ten of these peptides are common to both species of frog. Thirteen of the aurein peptides show wide-spectrum antibiotic and anticancer activity. Amongst the more active peptides are aurein 1.2 (GLFDIIKKIAESF-NH2), the smallest peptide from an anuran reported to have both antibiotic and anticancer activity; aurein 2.2 (GLLDIVKKVIGAFGSL-NH2) and aurein 3.1 (GLFDIVKKIAGHIAGSI-NH2). The aurein 4 and 5 peptides, e.g. aurein 4.1 (GLIQTIKEKLKELAGGLVTGIQS-OH) and aurein 5.1 (GLLDIVTGLLGNLIVDVLKPKTPAS-OH), show neither antibacterial nor anticancer activity.

Amino Acid Sequence↗
PubMed2000

Reciprocal interactions between perisynaptic Schwann cells and regenerating nerve terminals at the frog neuromuscular junction.

The perisynaptic Schwann cell (PSC) has gained recent attention with respect to its roles in synaptic function, remodeling, and regeneration at the vertebrate neuromuscular junction (NMJ). Here we test the hypothesis that, following nerve injury, processes extended by PSCs guide regenerating nerve terminals (NTs) in vivo, and that the extension of sprouts by PSCs is triggered by the arrival of regenerating NTs. Frog NMJs were double-stained with a fluorescent dye, FM4-64, for NTs, and fluorescein isothiocyanate (FITC)-tagged peanut agglutinin (PNA) for PSCs. Identified NMJs were imaged in vivo repeatedly for several months after nerve injury. PSCs sprouted profusely beginning 3-4 weeks after nerve transection and, as reinnervation progressed, regenerating NTs closely followed the preceding PSC sprouts, which could extend tens to hundreds of microns beyond the original synaptic site. The pattern of reinnervation was dictated by PSC sprouts, which could form novel routes joining neighboring junctions or develop into new myelinated axonal pathways. In contrast to mammals, profuse PSC sprouting in frog muscles was not seen in response to axotomy alone, and did not occur at chronically denervated NMJs. Instead, sprouting coincided with the arrival of regenerating NTs. Immunofluorescent staining revealed that in muscle undergoing reinnervation 4 weeks after axotomy, 91% of NMJs bore PSC sprouts, compared to only 6% of NMJs in muscle that was chronically denervated for 4 weeks. These results suggest that reciprocal interactions between regenerating NTs and PSCs govern the process of reinnervation at frog NMJs: regenerating NTs induce PSCs to sprout, and PSC sprouts, in turn, lead and guide the elaboration of NTs.

Animals↗
PubMed1977

Ultrastructure of traumatic cataractogenesis in the frog: a comparison with mouse and human lens.

Normal and needle-punctured lenses of Rana pipiens were examined with the electron microscope in order to characterize the sequence of ultrastructural changes that follow the injury over a 5-month period. Results were compared with those obtained previously in experimentally injured mouse and accidentally injured human lenses. The normal adult frog lens was found to have a morphology similar to that of mammalian lenses. As in the human, frog lens epithelial cells contained scattered microfilaments and were connected by desmosomes and gap junctions. They differed from mouse cells, which had been shown to lack desmosomes and to have microfilaments organized into dense bundles. These differences are postulated to be related to the degree of accommodative deformation of the lens displayed by these species. After injury, cellular debris and fibrin, accumulated in the wound, were phagocytized by extrinsic cells derived from the blood and ocular tissues. Leucocytes, pigmented cells and fibroblasts remained in the wound for eight weeks, along with epithelial cells which proliferated and migrated from the wound margins.Epithelial cells showed an increase in those organelles associated with protein synthesis and transport, and in microfilaments. In cataractous lenses, epithelial cells showed changes in matrix, and lens fibers became organized into smaller, denser compressed units. At five months, considerable healing had taken place, but localized opacities persisted in many frog lenses.

Animals↗
PubMed2001

Leaping lopsided: a review of the current hypotheses regarding etiologies of limb malformations in frogs.

Recent progress in the investigation of limb malformations in free-living frogs has underlined the wide range in the types of limb malformations and the apparent spatiotemporal clustering of their occurrence. Here, we review the current understanding of normal and abnormal vertebrate limb development and regeneration and discuss some of the molecular events that may bring about limb malformation. Consideration of the differences between limb development and regeneration in amphibians has led us to the hypothesis that some of the observed limb malformations come about through misdirected regeneration. We report the results of a pilot study that supports this hypothesis. In this study, the distal aspect of the right hindlimb buds of X. laevis tadpoles was amputated at the pre-foot paddle stage. The tadpoles were raised in water from a pond in Minnesota at which 7% of surveyed newly metamorphosed feral frogs had malformations. Six percent (6 of 100) of the right limbs of the tadpoles raised in pond water developed abnormally. One truncated right limb was the only malformation in the control group, which was raised in dechlorinated municipal water. All unamputated limbs developed normally in both groups. Three major factors under consideration for effecting the limb malformations are discussed. These factors include environmental chemicals (primarily agrichemicals), encysted larvae (metacercariae) of trematode parasites, and increased levels of ultraviolet light. Emphasis is placed on the necessary intersection of environmental stressors and developmental events to bring about the specific malformations that are observed in free-living frog populations.

Animals↗
PubMed1981

Ependymal astrocytes in the frog cerebellum.

We have examined the ependymal astrocytes of the frog cerebellar cortex in thin sections and freeze-fracture replicas. The somata border the fourth ventricle and give rise to basal processes whose daughter branches cross the molecular layer and terminate as subpial endfeet. Irregular lamellar appendages arise from the basal processes and their branches. In the molecular layer the appendages selectively ensheath apposed parallel fiber boutons and Purkinje cell dendritic spines. Other appendages ramify throughout the neuropil, some contributing to extensive pericapillary sheaths. Freeze-fractured ependymal astrocyte plasma membrane consistently has a greater concentration of intramembranous particles (IMPs) and IMPs of larger mean size than neuronal plasma membrane in the same replicas. Like the astrocytes of the mammalian central nervous system, frog ependymal astrocytes form numerous gap junctions with each other. However, orthogonal arrays of IMPs ("assemblies") were not observed. Ependymal cells in the frog cerebellum combine the morphology, and probably the functions, of both ependymal cells and astrocytes.

Animals↗