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PubMed2007

Germ cell depletion does not alter the morphogenesis of the fetal testis or ovary in the red-eared slider turtle (Trachemys scripta).

In the red-eared slider turtle, Trachemys scripta, both prospective male and female gonads contain primitive cord structures at the time when the gonad first forms. Primordial germ cells arrive in the gonad and accumulate on the coelomic surface. If testis development is initiated, these cords develop further at the same time that germ cells migrate from the coelomic surface and become sequestered in the interior of the cords. In contrast, in the developing ovary germ cells proliferate in a defined cortical domain, while the primitive cords regress and form flattened lacunae in the medulla. Because of their intimate association with these developmental processes, we investigated whether germ cells were required in turtles to establish the morphology of the fetal testis and ovary. We present evidence that normal morphological development of the fetal gonad occurs in both sexes in T. scripta after germ cell depletion, suggesting a conservation of developmental mechanisms across vertebrates.

Animals↗

PubMed1980

Ultrastructural morphology of the shell and shell membrane of eggs of common snapping turtles (Chelydra serpentina).

Common snapping turtles (Chelydra serpentina) lay nearly spherical, flexible-shelled eggs having an outer mineral layer composed of calcium carbonate in the aragonite form. The mineral layer is arranged into loosely organized groups of nodular shell units, with numerous spaces (or pores) between adjacent shell units. Shell units are structurally complex, consisting of an inner tip that is morphologically distinct from the main body of the shell unit. Contained within an intact shell unit at the interface of the tip and the main part of the shell unit is the central plaque, an apparent modification of the shell membrane that may serve to nucleate calcification of shell units during shell formation. The tips of shell units are firmly attached to a single, multilayered shell membrane throughout much of incubation. The calcareous layer begins to detach from the shell membrane about half-way through incubation, and changes in shell morphology attending this detachment indicate that snapping turtles may use the shell as a source of calcium during embryogenesis. The arrangement of the mineral layer into groups of shell units, the large number of spaces between shell units, and little or no interlocking of crystallites of adjacent shell units apparently are factors contributing to the ability of these eggs to swell as they absorb water.

Animals↗

PubMed1986

Morphology and histochemistry of the ambiens muscle of the red-eared turtle (Pseudemys scripta).

Six fiber types have been described in the ambiens muscle of red-eared turtles. These include one slow oxidative type, two fast oxidative types, two fast oxidative and glycolytic types, and one fast glycolytic type. Fiber types are non-randomly distributed throughout cross sections of the muscle. There is a decreasing gradient of oxidative staining and an increasing gradient of glycolytic staining along an axis from the superficial to deep regions of the muscle. The slow oxidative fibers are predominantly located within one or two fascicles of the superficial surface of the muscle. The fast glycolytic fibers are predominant in deep fascicles. In contrast to previous reports of histochemically monotypic intrafusal fibers in turtle muscle, ambiens muscle spindles have been observed containing one to eleven intrafusal fibers, including two fiber types. Fiber diameter and area are consistently smaller than observed in most extrafusal fibers. Spindles are predominantly located in superficial and cranial fascicles of the ambiens muscle and are located in regions characterized by extrafusal fibers with high oxidative activity.

Adenosine Triphosphatases↗

PubMed1995

Fiber-type composition of hindlimb muscles in the turtle, Pseudemys (Trachemys) scripta elegans.

A description is provided of the fiber-type composition of several hindlimb muscles of the adult turtle, Pseudemys (Trachemys) scripta elegans. In addition, cross-section areas of each fiber type and an estimation of the relative (weighted) cross-section area (wCSA) occupied by the different fiber types are also provided. Seven muscles were selected for study, based on their suitability for future neurophysiological analysis as components of the segmental motor system, and on their homologies with muscles in other vertebrates. The test muscles were iliofibularis (ILF), ambiens (AMB), external gastrocnemius (EG), extensor digitorum communis (EDC), flexor digitorum longus (FDL), tibialis anterior (TA), and peroneus anterior (PA). Serial sections of these muscles were stained for myosin adenosine triphosphatase (ATPase), NADH-diaphorase, and alpha-glycerophosphate dehydrogenase (alpha-GPDH), thereby enabling fiber-type classification on the basis of indirect markers for contraction speed and oxidative (aerobic) vs. glycolytic (anaerobic) metabolism. All muscles contained three fiber types: slow oxidative (SO; possibly including some non-twitch tonic fibers); fast oxidative glycolytic (FOG); and fast glycolytic (Fg). There were at least 30% FOG and 50% FOG + Fg fibers in the seven muscles, the extreme distributions being the predominantly glycolytic ILF vs. the predominantly oxidative FDL muscle (ILF--15.5% SO, 35.2% FOG, 49.3% Fg vs. FDL--49.1% SO, 41.1% FOG, 9.8% Fg). As in other species, the test muscles exhibited varying degrees of regional concentration (compartmentalization) of the different fiber types. This feature was most striking in ILF. Pronounced compartmentalization was also observed in AMB, EG, PA, TA, and EDC, whereas the distribution of fiber types in the highly oxidative FDL was homogeneous. In five of the seven muscles, fiber size was ranked with Fg > FOG > SO. In terms of wCSA, which provides a coarse-grain measure of the different fiber types' potential contribution to whole muscle peak force, all muscles exhibited a higher Fg and lower SO contribution to cross-section area than suggested by their corresponding fiber-type composition. The largest relative increase in wCSA vs. fiber-type composition were in the ILF and AMB muscles. We conclude that the turtle hindlimb provides some interesting possibilities for testing for a division of labor among different muscles during different movements (e.g., sustained vs. ballistic), and for study of the behavior of the different fiber (and motor unit) types under normal and perturbed conditions. The relationships between the present results and previous findings on homologous muscles of the mammalian (cat, rat) and reptilian (lizard) hindlimb are discussed.

Animals↗

PubMed2003

Role for calbindin-D28K in in vitro classical conditioning of abducens nerve responses in turtles.

Intracellular calcium has a pivotal role in synaptic modifications that may underlie learning and memory. The present study examined whether there were changes in immunoreactivity levels of the AMPA receptor subunits GluR2/3 and calcium binding proteins during classical conditioning recorded in the abducens nerve of in vitro brain stem preparations from turtles. The results showed that abducens motor neurons in unconditioned turtle brain stems were immunopositive for GluR2/3, calbindin-D28K, and calmodulin, but were immunonegative for parvalbumin. After classical conditioning, immunoreactivity for calbindin-D28K in the abducens motor nuclei was significantly reduced, whereas there were no significant changes in GluR2/3, calmodulin, or parvalbumin. This reduction in calbindin-D28K immunoreactivity was not observed following conditioning in the NMDA receptor antagonist AP-5, which blocked conditioned responses, suggesting that these changes are NMDA receptor-dependent. Moreover, the degree of the decrease in calbindin-D28K immunoreactivity was negatively correlated with the level of conditioning. Consistent with the immunocytochemical findings, Western blot analysis showed that calbindin-D28K protein levels were reduced after classical conditioning. The results support the hypothesis that in vitro classical conditioning of abducens nerve responses utilizes intracellular calcium-dependent signaling pathways that require NMDA receptor function and suggest a specific role for the calcium binding protein calbindin-D28K.

Abducens Nerve↗

PubMed1999

Effect of dichlorodiphenyltrichloroethane on sex determination of the common snapping turtle (Chelydra serpentina serpentina).

Recent evidence indicates that 1,1,1-trichloro-2, 2-bis(p-chlorophenyl)ethane (DDT) and some of its metabolites alter reproductive and endocrine function in wildlife. Exposure to such endocrine-disrupting compounds during embryonic development can affect sexual differentiation. The authors tested the hypothesis that dichlorodiphenyltrichloroethane (p,p'-DDE) causes feminization of the common snapping turtle (Chelydra s. serpentina), a species with temperature-dependent sex determination, during embryonic development. Eggs from eight clutches (total eggs tested=237) were incubated at a male-producing temperature (26 degrees C). At stage 14 of embryonic development, p,p'-DDE was applied topically at four concentrations and estrogen (estradiol-17 beta) was applied as a positive control. Although application of estrogen did induce female development at this temperature, application of p,p'-DDE did not affect sex determination at the exposure levels used. Residue analysis indicated that the amount of p,p'-DDE detected in the eggs 72 h after application was considerably less than the concentrations applied. However, the amounts that penetrated the shells were comparable to levels which have been found in moderately contaminated sites in the Great Lakes. These results indicate that p, p'-DDE, at levels that exist in the environment in the Great Lakes, does not cause the feminization of snapping turtles during embryonic development.

Animals↗

PubMed1994

Temperature-dependent sex determination in the snapping turtle: manipulation of the embryonic sex steroid environment.

In certain reptiles with temperature-dependent sex determination (TSD), estrogens act as a signal for female differentiation. Because aromatase produces estrogens from androgens, this enzyme plays a pivotal role in TSD. Whether androgens act as the signal for male differentiation in TSD species in not yet clear. We manipulated the hormonal environment in eggs of the common snapping turtle (Chelydra serpentina) to determine the effects of an estrogen (estradiol 17-beta), an aromatase inhibitor (fadrozole; CGS 16949A), and androgens (testosterone and dihydrotestosterone) on sex determination in this TSD species. Test solutions were applied topically to representative eggs (total tested = 1054 from 27 clutches) and incubated at two male-producing temperatures (24 and 26.5 degrees) and at a predominantly female-producing temperature (29 degrees). In this species, application of an estrogen induced female development at all temperatures tested. In contrast, the aromatase inhibitor had no effect at the male-producing temperatures, but induced male development at the predominantly female-producing temperature. At this temperature, aromatase inhibitor plus testosterone had a similar male-producing effect, but when applied alone, testosterone failed to augment male production. Dihydrotestosterone had a similar effect, in contrast to its reported androgenic effects in other TSD species. In the snapping turtle, male differentiation may not be androgen dependent; rather, it may proceed in the absence of female differentiation. In this species, female development is clearly estrogen-dependent and is altered by aromatase inhibition at female-producing temperatures. Our results not only provide additional evidence that sex steroids mediate gonadal differentiation in TSD species, but also suggest caution with respect to generalizations about the proximal mechanisms of TSD in reptiles.

Androgens↗

PubMed1994

Atrial natriuretic factor in the freshwater turtle Pseudemys scripta: a partial characterization.

The presence of natriuretic and vasorelaxant materials in the atria and ventricles of a Chelonian reptile, the freshwater turtle Pseudemys scripta, was verified and the active substance partially characterized. Crude atrial and ventricular extracts were acutely natriuretic and diuretic when administered to anesthetized rats increasing sodium excretion 23.5 +/- 7.9 and 5.11 +/- 18 microM Na/10 min/mg extract, respectively. Although atrial extracts were relatively more natriuretic than ventricular extracts, the total natriuretic content of ventricular extracts was approximately twofold greater. Moreover, partially purified atrial extracts were found to relax precontracted isolated rat aortic ring segments and exhibited immunoreactivity to mammalian ANP antisera. Also, the natriuretic and immunoreactive substance was purified further by size exclusion chromatography and a molecular weight of 3-5 kDa was determined. Using the bovine adrenal glomerulosa ANP receptor, this partially purified natriuretic substance displayed an apparent binding affinity similar to that of mammalian ANP. Finally, in order to demonstrate that this reptile is physiologically responsive to ANP, synthetic rat ANP (10 micrograms/kg) was shown to decrease arterial pressure in conscious turtles from a control value of 37.2 +/- 5.2 to 30.3 +/- 2.7 mm Hg. These data demonstrate that unlike other non-Chelonian reptiles, both the atria and the ventricles of this Chelonian reptile synthesize and store substantial levels of biologically active ANP-like materials, and further, that ANP is a highly conserved and primitive cardiac hormone.

Animals↗

PubMed1995

The relative effectiveness of androstenedione, testosterone, and estrone, precursors to estradiol, in sex reversal in the red-eared slider (Trachemys scripta), a turtle with temperature-dependent sex determination.

Androstenedione↗

PubMed1996

The relative effectiveness of estrone, estradiol-17 beta, and estriol in sex reversal in the red-eared slider (Trachemys scripta), a turtle with temperature-dependent sex determination.

In many turtles the temperature during the middle of incubation determines the gonadal sex of the hatchling. Sex steroid hormones have been implicated in temperature-dependent sex determination in the red-eared slider turtle, Trachemys scripta; nonaromatizable androgens are involved in male sex determination and estrogens and aromatizable androgens in female sex determination. Administration of exogenous estradiol-17 beta to eggs incubating at a temperature that normally produces only males can overcome the effect of temperature and result in all offspring being female. Further, estradiol-17 beta and incubation temperature synergize to produce a greater feminizing effect at intermediate incubation temperatures that produce mixed sex ratios. This study demonstrates that, in the red-eared slider, there is a complex interaction between incubation temperature, different estrogens, and the dosage effect of each hormone. There are changes in potency of different estrogens with incubation temperature such that estriol is more potent than estrone and estradiol-17 beta at 26 degrees (an all-male producing incubation temperature), estrone and estriol are equipotent to each other and more potent than estradiol-17 beta at 28.8 degrees (an incubation temperature that produced a male-biased sex ratio), and estradiol-17 beta is more potent than estrone and estriol at 29 degrees (an incubation temperature that produced equal numbers of males and females). These changes may be due to differences in synergism between the hormones and incubation temperature. Estriol treatment also resulted in cranially hypertrophied oviducts at all incubation temperatures in a dose-dependent manner, whereas animals treated with estradiol-17 beta and estrone had normal oviducts. These results support the hypothesis that estrogens are involved in the final common pathway of female sex determination in this species.

Animals↗

PubMed1997

Coexistence of alpha1 and beta adrenergic receptors in the liver of the frog Rana esculenta, the toad Bufo bufo, the lizard Podarcis sicula campestris, and the turtle Pseudemys picta elegans.

In mammals and birds the characteristics of alpha1 adrenergic receptors and their biological role in liver metabolism have been clearly described, although the predominance of receptor subtypes varies with species. In contrast, the actual presence of hepatic alpha1 adrenergic receptors in fish, amphibians, and reptiles has been questioned. Only recently has their existence been demonstrated in some fish species and also in the wood frog Rana sylvatica. The present study assessed the presence of alpha1 adrenergic binding sites on hepatic membranes of frogs, toads, lizards, and turtles using the specific alpha1 adrenergic receptor antagonist [3H]prazosin; for comparison, the binding of the specific beta adrenergic receptor antagonist [3H]CGP-12177A was evaluated in the same preparations. alpha1 Adrenergic receptors are indeed present in the liver of the ectotherms examined. Specific binding is saturable, reversible, and linear as a function of tissue concentration. The binding data indicated the presence of two classes of binding sites displaying high and low affinities with Kds in the nanomolar and micromolar ranges, respectively. The present study provides the first evidence for the presence of alpha1 adrenergic receptors in the liver of toad, lizard, and turtle while confirming their existence in another species of frog, Rana esculenta.

Adrenergic alpha-Antagonists↗

PubMed1998

Seasonal reproductive cycle of the Kemp's ridley sea turtle (Lepidochelys kempi).

The seasonal reproductive cycle of the Kemp's ridley sea turtle (Lepidochelys kempi) was studied under seminatural conditions at the Cayman Turtle Farm, Grand Cayman, British West Indies, from June 1987 to July 1988. Male L. kempi displayed a prenuptial rise in serum testosterone 4 to 5 months prior to the mating period (March). Male testosterone then declined sharply during the mating period. Female L. kempi also displayed a prenuptial rise in serum testosterone, estradiol, and total calcium 4 to 6 months prior to the mating period (March). Female testosterone and estradiol declined during the nesting period (April to July) immediately following the mating period (March). Elevated levels in female estradiol and total calcium corresponded with the period of vitellogenesis as determined from gel electrophoresis and ultrasonography. Serum thyroxine also fluctuated seasonally with elevated levels observed in females associated with the period of vitellogenesis. L. kempi displayed a distinct seasonal reproductive cycle in captivity. Nesting in the captive study group corresponded with nesting in the wild population at Rancho Nuevo, Mexico (April to July). Female endocrine cycles during the nesting period were similar to those observed in the wild population.

Animals↗

PubMed2000

Widespread expression of the mRNA encoding a novel vitamin D/thyroxine dual binding protein in the turtle Trachemys scripta.

Vitamin D-binding protein (DBP) is the major transport protein for the vitamin D sterols in a variety of mammalian and avian species. The DBP found in the emydid family of turtles is unique in that it exhibits high-affinity binding of both vitamin D and thyroxine (D/TBP). Sequence analysis has revealed that the emydid D/TBP is homologous to mammalian DBP and shares no homology with mammalian thyroxine-binding globulins. Northern blot analysis was used to examine the spatial profile of D/TBP transcription in the turtle Trachemys scripta. In both adults and hatchlings, two transcripts--the expected 1.5-kb full-length transcript and a second 0.8-kb transcript--were present in nearly all of the tissues examined. In adults, highest expression of the 1.5-kb transcript was seen in the kidney, gonad, and spleen, with lower levels in the liver and lung and no transcripts in skeletal muscle. In hatchlings, the full-length transcript was detected in a variety of tissues at similar levels. Injection of hatchlings with thyroxine increased levels of circulating D/TBP and transcript levels. These data are in marked contrast to observations in mammals in which transcription of DBP is confined predominantly to the liver. Further, the increase in circulating D/TBP associated with increased thyroidal activity may result from a direct or indirect activation of D/TBP transcription by thyroxine.

Aging↗

PubMed2000

Seasonal variation in reproductive steroids of male and female yellow-blotched map turtles, Graptemys flavimaculata.

From April to October of 1996 and 1997, we measured seasonal changes in plasma testosterone (T) in male and plasma T and estradiol-17beta (E(2)) in female yellow-blotched map turtles, Graptemys flavimaculata, from the Pascagoula River, Mississippi. In 1996, plasma from females was analyzed for progesterone. In 1997, ovaries of adult females were ultrasound imaged to determine stages of follicular development. Males exhibited peak T levels in September and October of both years, indicating fall gonadal activity. Males also exhibited a low level spring peak in T during April. Female E(2) levels increased significantly in May and June, the primary period of ovarian development. Females with preovulatory follicles did not have significantly higher E(2) levels than females with medium- or small-sized follicles. When compared to other freshwater turtle species, peak levels of E(2) were low. Testosterone levels did not follow a distinct yearly pattern in females. Progesterone levels were elevated in June when peak nesting was observed. Ultrasound and hormone data indicate that females lack ovarian development in the fall and produce on average only one clutch per year.

Animals↗

PubMed2000

cDNA cloning and functional expression of growth hormone receptor from soft-shelled turtle (Pelodiscus sinensis japonicus).

The growth hormone receptor (GHR) cDNA was cloned from the liver of soft-shelled turtle (Pelodiscus sinensis japonicus) using the polymerase chain reaction (PCR). Although GHR has been cloned from several mammalian and avian species, this is the first description of the reptilian receptor. As deduced from the nucleotide sequence, the precursor GHR of soft-shelled turtle (tGHR) is a protein of 615 amino acids which presents 72% identity with the chicken receptor and 57-64% identity with GHRs of several mammals. The tGHR expressed in COS-7 cells specifically bound human growth hormone (hGH) and was able to transduce an activation of transcription in the transfected cells. Binding of (125)I-hGH to the expressed receptor was decreased by the addition of excess unlabeled hGH, pig GH, and bream GH but not by pig insulin. The open reading frame of tGHR cDNA was inserted into the pSINrep/gfp (green fluorescence protein) vector and the tGHR-gfp fusion protein was stably expressed in baby hamster kidney (BHK) cells. Confocal imaging showed that tGHR-gfp was largely concentrated on the plasma membrane. Western blot analysis and deglycosylation treatment with PNGase F demonstrated that tGHR was a glycoprotein in BHK cells.

Amino Acid Sequence↗

PubMed1996

Mitochondrial gene order adjacent to the control region in crocodile, turtle, and tuatara.

We used the polymerase chain reaction and sequencing of mitochondrial gene junctions adjacent to the 5' end of the control region (light strand orientation) and the 3' end of ND6 to assess whether a representative crocodilian, turtle, or tuatara shares a unique mitochondrial gene order that is found in birds but not in mammals or amphibians. Turtles and crocodiles have the same gene order as mammals, except that crocodile has a tRNAPhe gene inserted between tRNAPro and the 5' end of the control region. Two different arrangements were detected at the 5' end of the control region in the tuatara, one resembling the mammalian (but with tRNAThr absent) and one resembling the avian gene order. Our data are consistent with the hypothesis that some tuatara mtDNAs within a single individual have undergone a deletion that removed the genes coding for cytochrome b and tRNAPro as well as 87 bp of the control region.

Alligators and Crocodiles↗

PubMed1997

Phylogenetic relationships of chelid turtles (Pleurodira: Chelidae) based on mitochondrial 12S rRNA gene sequence variation.

Conflicting phylogenies have been proposed for the Chelidae (Testudines: Pleurodira), a family of side-necked turtles found only in Australasia and South America. Sequence data from the mitochondrial 12S rRNA gene were used to test these phylogenies. In total, 411 nucleotides were sequenced for each of 16 chelid species, including all 11 recognized chelid genera and, as outgroups, 5 genera of Pelomedusidae (Testudines: Pleurodira). Analyses using parsimony and neighbor joining algorithms strongly support the division of Australian Chelidae into the three monophyletic groups initially suggested by Burbidge et al. (1974; Copeia 2: 392-409): Chelodina (bootstrap value 99%), the Emydura group (87%), and Pseudemydura. The analyses suggest that the Australian chelids are a monophyletic lineage (64%), with the Australian long-necked turtles, Chelodina, more closely related to the Australian short-necked chelids than to the long-necked South American species. These relationships are in contrast to those of Gaffney (1977; Am. Mus. Novitates 2620: 1-28). The species of Australian long-necked chelids consistently form a monophyletic clade, with Chelodina longicollis and Chelodina oblonga as sister taxa. The data failed to resolve relationships among the Australian short-necked taxa: Emydura, the Elseya latisternum group, the Elseya dentata group, Rheodytes, and Elusor. Unlike Gaffney (1977), we find some weak support (58%) for Pseudemydura as the closest relative of the other Australian short-necked taxa. With the exception of Hydromedusa, the South American taxa are monophyletic and the subgenera of Phrynops are paraphyletic.

Animals↗

PubMed1993

Immunocytochemical localization of corticotropin-releasing factor in the brain of the turtle, Mauremys caspica.

Brain sections of the turtle, Mauremys caspica were studied by means of an antiserum against rat corticotropin-releasing factor. Immunoreactive neurons were identified in telencephalic, diencephalic and mesencephalic areas such as the cortex, nucleus caudatus, nucleus accumbens, amygdala, subfornical organ, paraventricular nucleus, hypothalamic dorsolateral aggregation, nucleus of the paraventricular organ, infundibular nucleus, pretectal nucleus, periventricular grey, reticular formation and nucleus of the raphe. Many immunoreactive cells located near the ependyma were bipolar, having an apical dendrite that contacted the cerebrospinal fluid. Immunoreactive fibers were seen in these locations and in the lamina terminalis, lateral forebrain bundle, supraoptic nucleus, median eminence, neurohypophysis, tectum opticum, torus semicircularis and deep mesencephalic nucleus. Parvocellular bipolar immunoreactive neurons from the paraventricular and infundibular nuclei projected axons that joined the hypothalamo-hypophysial tract and reached the outer zone of median eminence, and the neural lobe of the hypophysis where immunoreactive fibers terminated close to intermediate lobe cells. From these results it can be concluded that, as in other vertebrates, corticotropin-releasing factor in the turtle may act as a releasing factor and, centrally, as a neurotransmitter or neuromodulator.

Animals↗