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PubMed1999

The effect of isovolemic anaemia on blood O2 affinity and red cell triphosphate concentrations in the painted turtle (Chrysemys picta).

The blood oxygen affinity of vertebrates is regulated, in part, through changes in red cell phosphate levels and increased oxygen affinity during reductions in inspired oxygen and is a well-described and common feature. However, during anaemia, when oxygen delivery is compromised by a reduction in blood oxygen carrying capacity, a lowering of blood oxygen affinity will facilitate oxygen unloading in the tissues, while oxygen loading at the gas exchange organ is not impaired. The present study investigated the effects of artificially induced anaemia in vivo on the blood oxygen affinity and red cell nucleoside triphosphate (NTP) concentrations in the turtle, Chrysemys picta. Blood was obtained from conscious animals through an arterial catheter and oxygen equilibrium curves were determined using the Tucker method while NTP concentrations were analyzed spectrophotometrically. Before induction of anaemia haematocrit averaged 23% and P50 was 18.5 +/- 0.7 with a NTP/Hb of 0.20 +/- 0.01 (mmol/mmol). After the haematocrit had been reduced to approximately 10% by bleeding (48-96 h) (blood volume was maintained by re-infusion of plasma and Ringer) there were no effects on P50 or red cell NTP concentrations. Thus, in contrast to fish and mammals, turtles do not exhibit a change in blood oxygen affinity during anaemia.

Anemia↗
PubMed1999

Distribution of glycogen phosphorylase and cytochrome oxidase in the central nervous system of the turtle Trachemys dorbigni.

Glycogen phosphorylase (GP) and cytochrome oxidase (CO) activities were mapped histochemically in the brain of the turtle Trachemys dorbigni. In the telencephalon, both activities occurred in the olfactory bulb, in all cortical areas, in the dorsal ventricular ridge, striatum, primordium hippocampi and olfactory tubercle. In the diencephalon, they were identified in some areas of the hypothalamus, and in rotundus and geniculate nuclei. Both reactions were detected in the oculomotor, trochlear, mesencephalic trigeminal nuclei, the nucleus of the posterior commissure, torus semicircularis, substantia nigra and ruber and isthmic nuclei of the mesencephalon. In all layers of the optic tectum GP activity was found, but CO only labelled the stratum griseum centrale. In the medulla oblonga both enzymes appear in the reticular, raphe and vestibular nuclei, locus coeruleus and nuclei of cranial nerves. In the cerebellum, the granular and molecular layers, and the deep cerebellar nuclei were positive for both enzymes. The Purkinje cells were only reactive for CO. In the spinal cord, motor and commissural neurones exhibited a positive reaction for the two enzymes. However, CO also occurred in the marginal nucleus and in the lateral funiculus. These results may be useful as a basis for subsequent studies on turtle brain metabolism.

Animals↗
PubMed1997

Long-term recording of olfactory and vomeronasal stimulant-induced waves from the turtle main olfactory bulb and accessory olfactory bulb.

Recording of stimulant-induced waves (bulbar responses) from the main olfactory bulb is a useful tool for measuring quantitative stable olfactory responses. There is a good relationship between the olfactory bulbar response, olfactory nerve response and electroolfactogram (EOG), suggesting that the bulbar response reflects events in receptor cells. The modern whole-cell recording technique offers direct information on olfactory transduction in single cells, but it requires long experimental periods and many animals. On the other hand, analysis of bulbar responses provides useful information and requires the use of few animals. For example, we found that cAMP-increasing and IP3-increasing odorants were not distinctly received by the turtle olfactory organ by measuring olfactory bulbar responses and analyzed with a multidimensional scaling from about 60 animals. However, to record similar odor responses from isolated turtle olfactory neurons, at least 200 animals would be necessary. Bulbar responses are recorded with electrodes implanted into or located on the main olfactory bulb. When electrodes are located on the olfactory bulb surface, it is possible to record stable responses over a period of 3 days. These methods were applied successfully to the accessory olfactory bulb. In this paper, we describe the protocols used for recording of the stimulant-induced waves from the main and accessory olfactory bulb.

Adaptation, Physiological↗
PubMed1999

Origin and propagation of spontaneous electrographic sharp waves in the in vitro turtle brain: a model of neuronal synchronization.

OBJECTIVES: Neuronal synchronization is a basic feature in the generation of epileptiform discharges. Spontaneous large sharp waves (LSWs) can be recorded in the turtle brain in vitro, indicating the synchronous activation of large neuronal populations. The aim of this study was to analyze the spatial and temporal distribution of LSWs within the brain; the participation of glutamate in LSWs generation was also investigated. METHODS: Extracellular field potentials were recorded in vivo (n = 4) and in vitro (n = 36). LSWs were recorded from cerebral cortex, optic tectum, and thalamus. RESULTS: LSWs were recorded from cerebral cortex, optic tectum and thalamus. No LSWs were observed in cerebellum and brain stem. In some experiments, LSWs could be recorded only from medial cortex. Latency studies demonstrated that, within each hemisphere, medial cortex led the generation of LSWs; in addition, isolated medial cortex could sustain LSWs. Intracortical laminar field potentials in medial cortex indicated that LSWs generate mainly in the molecular layer, probably at pyramidal cell dendrites. Pharmacological experiments demonstrated that NMDA and non-NMDA glutamate receptors are involved in LWSs generation. CONCLUSIONS: These results suggest that turtle medial cortex is the pacemaker area for LSWs generation and it can be a useful model to study cellular and circuital mechanisms of neuronal synchronization.

Animals↗
PubMed1988

Neural organization of the retina of the turtle Mauremys caspica: a light microscope and Golgi study.

The organization of the retina of the turtle species Mauremys caspica, found in fresh water ponds of Israel, has been examined by light microscopical techniques including examination of fresh wholemount retina, one micron blue-stained vertical sections and Golgi-stained material. The anatomical findings on Mauremys retina have been compared with those of the Pseudemys retina (Kolb, 1982) which is more commonly used for electrophysiological and neurochemical studies in the USA. The photoreceptors of Mauremys are similar in type and oil droplet content to Pseudemys photoreceptors except for the double cone in Mauremys. This cone type appears more abundant than in Pseudemys and the principal member contains a yellow oil droplet instead of an orange oil droplet. Golgi staining reveals that the cell types that have been seen in Pseudemys are found in Mauremys with identical morphology. In addition, two amacrine cell types that were not before described for Pseudemys have been added to the classification. One of these is the tristratified dopaminergic amacrine cell described in immunocytochemical studies (Witkovsky et al., 1984; Nguyen-Legros et al., 1985; Kolb et al., 1987). We have used these anatomical studies on Pseudemys and Mauremys retina to form a catalogue of neural types for the turtle retina in general. We conclude with an attempt to combine findings from anatomy, electrophysiology, and neurochemistry to form an overview of the organization of this reptilian retina.

Animals↗
PubMed1991

Spatial properties of ganglion cell activity in the turtle retina.

Ganglion cell activity in response to drifting or alternating stimulus patterns was recorded in an eyecup preparation of the turtle. The stimuli were sinusoidal gratings with various spatial frequencies and spatial phases. Spike histograms of the activity were Fourier-analyzed. The results indicated that the retinal circuitry always functioned in a nonlinear manner. The average, first, and second harmonics of the activity were all a function of stimulus conditions, including spatial phase. Similar outcomes were obtained with full-field (retina-wide) and a small-field (1 mm diameter) stimulation. In the middle of the spatial-frequency range, the ratio of second to first harmonic activity was higher in response to static, contrast-reversing gratings than for drifting, constant-contrast ones. The findings suggest that, as distinct from the usual X, Y, and W segregation of ganglion cell properties in other retinas, there is a single class of cells whose responses seem to have attributes of one or another of the usual categories depending on stimulus characteristics. This may be the result of a degree of stimulus-dependent functional modification in the retinal input circuitry to the turtle ganglion cells.

Animals↗
PubMed1989

Immunocytochemical localization of glycine in the retina of the turtle (Pseudemys scripta).

We have localized glycine-like immunoreactivity to provide new anatomical detail about glycinergic neurons in the turtle retina. A rabbit antiserum directed against a glycine/albumin conjugate was used with standard fluorescent and avidin-biotin labeling techniques. Some processes in the outer plexiform layer and many processes in the inner plexiform layer, numerous somata in the inner nuclear layer, and isolated somata in the ganglion cell layer were immunoreactive. The vast majority of labeled neurons were amacrine cells. One class of amacrine cells had well-labeled somata near the inner nuclear/inner plexiform layer border, which gave rise to thick primary processes that entered the inner plexiform layer and arborized near the border of strata 1 and 2 and in stratum 3. A second class of glycinergic neurons, consisting of putative interplexiform cells, was unique in that it gave rise to dendritic arborizations in both the outer plexiform layer and the inner plexiform layer. Some of the immunoreactive neurons in the ganglion cell layer were apparently displaced amacrine cells, while others were probably true ganglion cells because they gave rise to labeled axons, and many labeled axons were visible in the ganglion cell axon layer. These results suggested that glycine played an extensive role in the turtle retina, and that it was involved in many diverse synaptic interactions in both the outer plexiform layer and the inner plexiform layer.

Animals↗
PubMed1990

Effects of calcium ions on L-type horizontal cells in the isolated turtle retina.

A technique by which the retina can be isolated from the turtle eye is described. Scanning electron microscopy revealed morphological variability between preparations and also between regions of the same one. Large areas were often totally free of any pigment epithelial cells, yet contained a high proportion of photoreceptors with complete outer segments. However, adjacent regions may contain photoreceptors without outer segments or with fragmented ones. The physiological properties of the horizontal cells also demonstrated large variability between different preparations. In all cases, lowering calcium concentration from 2 mM to 0.1-0.5 mM depolarized the horizontal cells and augmented the amplitude of the maximum photoresponses. However, these effects were accompanied by changes in the photoresponse kinetics and by a reduction in the horizontal cell sensitivity to light. Moreover, prolonged exposure to low calcium induced permanent damage to the retina as was indicated by the reduction in the response amplitude after superfusion with 2 mM calcium solution had been resumed. The toxic effects of low calcium were most apparent when superfusion with 0.1-1.0 microM calcium concentration was performed. These solutions induced complex time-dependent effects on the resting potential of horizontal cells and on the amplitude and kinetics of the photoresponses. We conclude from these observations that the normal concentration of extracellular calcium in the turtle retina is in the 2 mM range.

Animals↗
PubMed1989

Signal integration at the pedicle of turtle cone photoreceptors: an anatomical and electrophysiological study.

The morphology of the axon which connects the cell body and pedicle of turtle cone photoreceptors was studied by light and electron microscopy. The axon which contains numerous synaptic vesicles, some endoplasmic reticulum, and a few cisternae is basically filled with cytoplasm. The length of the axon is related to the class of cone and varies slightly with retinal location, with axons as short as 3-6 microns found in red cones, and as long as 60 microns in cones containing colorless oil droplets. By simultaneously voltage clamping the cell body and pedicle regions of single isolated cones, we measured the longitudinal axonal resistance and the cell body and pedicle membrane resistances. For each cell studied, the axonal resistance of cones with short axons was lower than the cell and pedicle membrane resistances. Thus, the cell can be considered to be an isopotential structure. However, in some cones with long axons, the axonal, cell body, and pedicle resistances were comparable. The pedicles of these cones, therefore, could act like summing points and may provide a locus for spatial signal integration. Electrical coupling between the principal and accessory members of double cones was also studied. Electron-microscopic observation of the membrane junction between the apposed inner segments of the double cones in the intact retina show narrow segments which resemble gap junctions. However, in every double cone studied in culture, passing currents into one member of the double cone did not result in measurable current flow in the adjacent cell. Thus, the two members of the double cone, isolated from the turtle retina, are not electrically coupled.

Animals↗
PubMed1990

Wavelength-dependent temporal properties of retinal horizontal cells in turtles.

Electrical responses of luminosity horizontal cells (L cells) to monochromatic stimuli were analyzed by intracellular recordings in the retinas of the freshwater turtle (Pseudemys scripta elegans) and of the sea turtle (Chelonia mydas mydas). Light intensity, duration, and wavelength were varied to assess temporal effects. For a given intensity of monochromatic light, response amplitude increased with stimulus duration until maximum amplitude occurred at a specific duration. This suprathreshold metric of temporal integration is called here summation time, and it is wavelength-dependent. L cells always had some level of red-sensitive cone input, although in some cells inputs from green- and blue-sensitive cones were also observed. For these latter cells, summation times were shorter for 640-nm than for 540-nm or 450-nm lights. These results were most evident in cells that received dominant inputs from blue- or green-sensitive cones. Responses of some other L cells were almost completely dominated by inputs from red-sensitive cones. Summation times of these cells were not wavelength-dependent. But when these inputs also included green-sensitive cones, shorter summation times were obtained to 640-nm light than to 540-nm light, even though dominant inputs were still from red-sensitive cones. These results, obtained from both retinal and 3,4-dehydroretinal photopigment systems, are consistent with reported observations in Pseudemys scripta elegans that show linear responses of red-sensitive cones to have shorter integration times and times-to-peak than green-sensitive cones. Responses from horizontal cells dominated by blue-sensitive cone inputs were the most sensitive of all; they also had the longest summation times.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMedDate not supplied

Synaptic connections involving immunoreactive glycine receptors in the turtle retina.

The distribution of glycine receptors in the turtle retina was studied with the aid of a monoclonal antibody that detects the 93-kD protein associated with the strychnine-sensitive glycine receptor. Light microscopically, receptors were found in the inner plexiform layer and, more sparsely, in the innermost parts of the inner nuclear layer. No receptors were seen to be associated with photoreceptor cells, horizontal cells, or any other structures in the distal inner nuclear layer or outer plexiform layer. Ultrastructurally, glycine receptors were found on the inner face of postsynaptic membranes of processes from amacrine and presumed ganglion cells and always involved amacrine cell processes as the presynaptic element. Such glycine receptor immunoreactive synapses onto amacrine cell processes were distributed throughout the inner plexiform layer with a peak density near the middle. On the other hand, output synapses onto ganglion cell processes displaying immunoreactive glycine receptor sites showed a bimodal distribution in the inner plexiform layer. Glycine receptor immunoreactivity was not detected on bipolar cells, but presumed glycine-utilizing processes (i.e. those presynaptic to immunoreactive glycine receptors) were occasionally found to be postsynaptic in bipolar cell dyads. The majority of the synaptic input to the presumed glycine-utilizing amacrine cell processes was from other amacrine processes, some of which were themselves glycine utilizing. The observations suggest that glycinergic synapses in the turtle retina are, to a large extent, engaged in processing interamacrine signals.

Animals↗
PubMedDate not supplied

A model for optokinetic eye movements in turtles that incorporates properties of retinal-slip neurons.

The turtle's optokinetic response is described by a simple model that incorporates visual-response properties of neurons in the pretectum and accessory optic system. Using data from neuronal and eye-movement recordings that have been previously published, the model was realized using algebraic-block simulation software. It was found that the optokinetic response, modelled as a simple negative feedback system, was similar to that measured from a behaving animal. Because the responses of retinal-slip detecting neurons corresponded to the nonlinear, closed-loop optokinetic response, it was concluded that the visual signals encoded in these neurons could provide sufficient sensory information to drive the optokinetic reflex. Furthermore, it appears that the low gain of optokinetic eye movements in turtles, which have a negligible velocity storage time constant, may allow stable oculomotor output in spite of neuronal delays in the reflex pathway. This model illustrates how visual neurons in the pretectum and accessory optic system can contribute to visually guided eye movements.

Animals↗
PubMedDate not supplied

Is the input to a GABAergic synapse the sole asymmetry in turtle's retinal directional selectivity?

We examined the effects of picrotoxin and pentylenetetrazol (PTZ) on the responses to motions of ON-OFF directionally selective (DS) ganglion cells of the turtle's retina. These drugs are antagonists of the inhibitory neurotransmitter GABA. For continuous motions, picrotoxin markedly reduced the overall directionality of the cells. In 21% of the cells, directional selectivity was lost regardless of speed and contrast. However, other cells maintained their preferred direction despite saturating concentrations of picrotoxin. And in most cells, loss, maintenance, or even reversal of preferred and null directions could occur as speed and contrast were modulated. In 50% of the cells, reversal of preferred and null directions occurred at some condition of visual stimuli. However, picrotoxin did not tend to alter the preferred-null axis for directional selectivity. For apparent motions, picrotoxin made motion facilitation, which normally occurs exclusively in preferred-direction responses, to become erratic and often occur during null-direction motions. Finally, PTZ had effects similar to picrotoxin but with less potency. The results in this paper indicated that models of directional selectivity based solely on a GABAergic implementation of Barlow and Levick's asymmetric-inhibition model do not apply to the turtle retina. Alternative models may comprise multiple directional mechanisms and/or a symmetric inhibitory one, but not asymmetric facilitation.

Afferent Pathways↗
PubMed1991

Synaptic inputs to physiologically defined turtle retinal ganglion cells.

Two physiologically distinct, HRP-marked turtle retinal ganglion cells were examined for their morphology, GABAergic, glycinergic, and bipolar cell synaptic inputs, using electron-microscopic autoradiography and postembedding immunocytochemistry. One cell was a color-opponent, transient ON/OFF ganglion cell. Its center response to red was a sustained hyperpolarization, and its center response to green was a depolarization with increased spiking at onset. The HRP-injected cell most resembled G6, from previous Golgi-impregnation studies (Kolb, 1982; Kolb et al., 1988). It was a narrow-field bistratified cell, whose two broad dendritic strata peaked at approximately levels L20-25 (sublamina a) and L60 (sublamina b) of the inner plexiform layer. Bipolar cell synapses onto G6 were found evenly distributed between its distal and proximal dendritic strata, spanning L20-75. These inputs probably originated from several different bipolar cells, reflecting the complexity of the center response. GABAergic inputs were found onto both the distal and proximal strata, from near L20-L85. Only a few glycinergic inputs, confined to dendrites at L50-70, were observed. A second ganglion cell type that we physiologically characterized and HRP-injected had sustained ON-center, sustained OFF-surround responses. Two examples were studied; both were bistratified in sublamina b, near L60-70 and L85-100, with branches up to near L40. They resembled G10, from previous Golgi-impregnation studies (Kolb, 1982; Kolb et al., 1988). One cell was partially reconstructed to look at the distributions of GABAergic and glycinergic amacrine cell, and bipolar cell inputs. Although synapses from bipolar cells were equally divided between the two major dendritic strata of G10, the inputs to the distal stratum were close to the soma, and the inputs to the more proximal stratum were on the peripheral dendrites. This arrangement may reflect input from two distinct types of ON-bipolar cell. GABAergic and glycinergic inputs to G10 costratified to both strata and to the distal branches; but where glycinergic inputs were found distributed throughout the arbor, GABAergic inputs appeared to be confined to peripheral dendrites. We hypothesize on the neural elements involved and the circuitry that may underlie the physiologically recorded receptive fields of these two very different ganglion cell types in the turtle retina.

Animals↗
PubMed1991

Effects of synaptic drugs on turtle optokinetic nystagmus and the spike responses of the basal optic nucleus.

Behavioral and electrophysiological measures were used to elucidate the retinal modulation of oculomotor control in the turtle. Eye movements were recorded following intravitreal applications of 2-amino-4-phosphonobutyrate (APB) and the GABA antagonists picrotoxin and bicuculline. Visual responses of single basal optic nucleus (BON) neurons of the accessory optic system were studied in parallel experiments. The effectiveness of APB, a glutamate analog thought to act selectively on the retinal ON pathway, was assessed independently by recording electroretinograms or ganglion cell activity. Injections of APB into the turtle's eye reduced or blocked the injected eye's ability to drive horizontal optokinetic nystagmus, as also observed in rabbit and cat (Knapp et al., 1988; Yucel et al., 1989). Single-unit recordings from the BON during APB superfusion (50-200 microM APB) of the contralateral retina demonstrated that these cells, which are direction-sensitive and respond to the offset of light flashes, have their responses to moving stimuli blocked by APB. During the APB effect, GABA antagonists were applied to the same eye. Although moderate doses of APB were sufficient to block optokinetic or BON light responses, the addition of GABA blockers still elicited a spontaneous temporal-to-nasal nystagmus (Ariel, 1989) or visually responsive yet direction-insensitive responses from BON cells (Schuerger et al., 1990). These results are discussed in terms of the retinal output to pathways involved in oculomotor control of optokinetic nystagmus.

Aminobutyrates↗
PubMedDate not supplied

Modulation of endogenous dopamine release in the turtle retina: effects of light, calcium, and neurotransmitters.

In the turtle retina, dopamine has been observed in a small population of amacrine cells. Whereas the effect of dopamine has been intensively studied, knowledge about the release of this transmitter and the neuronal control of its release are still poorly understood. We therefore decided to study the release of endogenous dopamine. Isolated retinas were superfused with Ringer's solutions and stimulated with increased potassium, light, or drugs which interfere with neurotransmitter systems. Dopamine was analyzed by using aluminum-oxide extraction and high-pressure liquid chromatography (HPLC) with electrochemical detection. Increased potassium (25 mM) caused a five-fold increase in the basal release. When calcium was replaced by cobalt, no increase was induced by 25 mM potassium. Flickering light increased the basal release of endogenous dopamine by a factor of three. The effect of flickering light was greater in the presence of additional steady background illumination. Kainate (10 microM), an agonist for excitatory amino acids, doubled the basal dopamine release. Bicuculline (10 microM), a gamma-amino butyric acid (GABA) antagonist, increased the release to about six times the basal level. Naloxone (10 microM), an opiate antagonist, increased the release to eight times the basal level. These findings suggest that dopamine is released from amacrine cells in the turtle retina in a calcium-dependent manner, which is most likely a vesicular release. Dopamine release is induced by flickering light vs. darkness and vs. steady background illumination. A moderate background illumination alone does not significantly increase basal dopamine release.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1991

Refractive state, contrast sensitivity, and resolution in the freshwater turtle, Pseudemys scripta elegans, determined by tectal visual-evoked potentials.

Visual-evoked potentials (VEPs) were recorded from the surface of the optic tectum of the freshwater turtle, Pseudemys scripta elegans, in response to phase reversal of square-wave gratings of different spatial frequency and contrast. The refractive state of a group of 12 turtles in air was assessed from VEPs by placing trial lenses in front of the eye. The group mean refraction did not differ significantly from emmetropia, as compared to 4.8 diopters of hyperopia when refracted retinoscopically. The difference was explained by the retinoscopic reflex originating from the interface between vitreous humor and retina. Peak VEP amplitude was approximately linear with log grating contrast; extrapolation to zero VEP amplitude yielded contrast thresholds as low as 1%. High spatial-frequency cutoffs ranged from 4.4-9.9 cycle/deg in different animals, the highest values corresponding to the intercone spacing in the area centralis and to behavioral measures of acuity in a related species.

Adaptation, Ocular↗
PubMed1991

Ocular dimensions and schematic eyes of freshwater and sea turtles.

Measurements were made of the ocular dimensions from living and frozen eyes of one species of freshwater turtle, Pseudemys scripta elegans, and of three species of marine turtles, Chelonia mydas, Dermochelys cariacea, and Eretmochelys imbricata. Estimates of refractive error by retinoscopy were also obtained with eyes in air and under water. The results suggest that unaccommodated eyes of all four species are approximately emmetropic in air but strongly hyperopic in water. Schematic eyes were calculated for each species in both air and water.

Adaptation, Physiological↗