Synaptic responses of medullary neurons of turtles to stimulation of the locomotor region

Responses of medullary neurons to microstimulation of the locomotor region by a current of up to 30 µA were studied by intracellular recording in turtles. The resting potential recorded in these neurons was from 22 to 42 mV. Depolarization PSPs (EPSPs) were recorded in 43 neurons, hyperpolarization PSPs (IPSPs) in 12, and mixed in 36. Synaptic discharges were observed in 29 neurons. Of these cells 11 generated action potentials without visible PSPs. The latent period of the PSPs was most frequently between 2 and 8 msec. The time from the beginning of the EPSP to the beginning of the action potential was 1–3 msec if the response index was high, but in the case of weaker stimulation, it began to fluctuate strongly and lengthened. Unitary EPSPs were recorded in 15 neurons and IPSPs in three. Their amplitude was 0.6–0.8 mV, from 2 to 12 times less than the depolarization threshold (1–7 mV). These results, together with those obtained previously by extracellular recording of medullary unit activity in turtles and cats, are used to discuss the possible mechanism of spread of locomotor activity.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 14, No. 2, pp. 122–129, March–April, 1982.     

The rhombencephalic recess in the rat

Summary The rhombencephalic recess, an ependymal organ, has been studied for the first time by light- and electron microscopy. It is situated mediosagittally on the floor of the rhomboid fossa at the level of the colliculus facialis. The recess and the superimposed tissue are built up by tanycytes, their apices being connected by tight junctions. HRP, injected into the c.s.f., does not penetrate into the intercellular clefts of the recess area. The recess area reveals a certain autonomy regarding its supply with arteries and capillaries. A bloodbrain barrier exists, but shows slight leakage in circumscribed areas as a result of intense transendothelial vesicular transport. The organization of the recess area is compared with that of other ependymal organs, especially circumventricular organs.The skilful technical assistance of Miss K. Bielenberg, Mrs. H. Prien, Mrs. E. Schöngarth and Mrs. H. Schöning is thankfully acknowledgedSupported by the Deutsche Forschungsgemeinschaft (Kr 569/1 and SFB 34/D4) and Stiftung Volkswagenwerk

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The effects of temperature and inter-individual variation on the locomotor performance of juvenile turtles

The effects of temperature on aquatic and terrestrial locomotor performance, including measures of burst speed, endurance, and righting response, the inter-individual correlation between measures of locomotor performance, and the temporal repeatability of performance were assessed in juvenile western painted turtles, Chrysemys picta bellii. Locomotor performance increased as temperature increased, with Q ₁₀ values ranging from 1.33 to 1.98 for burst speed and 2.28 to 2.76 for endurance measures. Righting response performance also increased with temperature. Aquatic and terrestrial measures of locomotor performance were highly correlated; however, righting response was not correlated with any other measure of performance. Measures of terrestrial locomotor performance were highly repeatable over the entire 30-week study period, whereas aquatic locomotor performance was only repeatable through week 12. The righting response was repeatable over a 6-week study period. Both the interindividual variation and temperature effects on locomotor performance likely influences the survival of turtles, especially juveniles, by affecting the length of time turtles are exposed to potential predators and their ability to escape.     

Forebrain projections to the hypothalamic locomotor region in cats

Descending projections from the cortex and basal ganglia to the hypothalamic locomotor region were studied in cats by the retrograde axonal transport of horse-radish peroxidase method. Neurons forming direct projections to the physiologically identified hypothalamic locomotor region are diffusely scattered over different gyri, but predominantly in areas 4 and 6 of the motor cortex, and also in the entopeduncular nucleus. Powerful cortico- and pallido-hypothalamic projections mainly do not reach as far as the most caudal zones of the hypothalamus, where the region whose electrical stimulation evokes locomotion lies.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 2, pp. 255–263, March–April, 1985.     

Functional diversity for body actions in the mesencephalic locomotor region

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Testing for evolutionary trade-offs in a phylogenetic context: ecological diversification and evolution of locomotor performance in emydid turtles

The evolution of ecological trade-offs is an important component of ecological specialization and adaptive radiation. However, the pattern that would show that evolutionary trade-offs have occurred between traits among species has not been clearly defined. In this paper, we propose a phylogeny-based definition of an evolutionary trade-off, and apply it to an analysis of the evolution of trade-offs in locomotor performance in emydid turtles. We quantified aquatic and terrestrial speed and endurance for up to 16 species, including aquatic, semi-terrestrial and terrestrial emydids. Emydid phylogeny was reconstructed from morphological characters and nuclear and mitochondrial DNA sequences. Surprisingly, we find that there have been no trade-offs in aquatic and terrestrial speed among species. Instead, specialization to aquatic and terrestrial habitats seems to have involved trade-offs in speed and endurance. Given that trade-offs between speed and endurance may be widespread, they may underlie specialization to different habitats in many other groups.     

Connections of the subthalamic and mesencephalic "locomotor regions" in rats

Responses of single neurons to stimulation of the subthalamic "locomotor region" were recorded extracellularly in the tegmentum mesencephali of rats. Latent periods of response of different neurons varied from 1 to 13 msec. The thresholds of the unit responses usually did not exceed the threshold for eliciting locomotion. The results are evidence of direct and oligosynaptic connections between the "locomotor regions" of the subthalamus and mesencephalon.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 275–280, May–June, 1977.     

Electrocardiography in turtles

The normal ECG of turtles is similar to that of man, but low voltage waves are the rule. The Q and S waves are uncommon. Sex differences are described. In the female the P-R interval is shorter, but the R and T waves are longer and have a lower voltage Unfed males and females show occasional arrhythmias and marked electrical differences from the normal in the voltages and durations of the ECG parameters, but the sex differences which characterize the fed turtles are slightly reduced. Several significant differences are described in the ECG in the surgical stage of anesthesia produced by ether or by sodium pentobarbital. Species differences are observable in the ECG records, but these differences may reside largely in the different pacemaker rates.     

The ecology of overwintering among turtles: where turtles overwinter and its consequences

Turtles are a small taxon that has nevertheless attracted much attention from biologists for centuries. However, a major portion of their life cycle has received relatively little attention until recently - namely what turtles are doing, and how they are doing it, during the winter. In the northern parts of their ranges in North America, turtles may spend more than half of their lives in an overwintering state. In this review, I emphasise the ecological aspects of overwintering among turtles, and consider how overwintering stresses affect the physiology, behaviour, distributions, and life histories of various species.Sea turtles are the only group of turtles that migrate extensively, and can therefore avoid northern winters. Nevertheless, each year a number of turtles, largely juveniles, are killed when trapped by cold fronts before they move to safer waters. Evidently this risk is an acceptable trade-off for the benefits to a population of inhabiting northern developmental habitats during the summer.Terrestrial turtles pass the winter underground, either in burrows that they excavate or that are preformed. These refugia must provide protection against desiccation and lethal freezing levels. Some burrows are extensive (tortoise genus Gopherus), while others are shallow, or the turtles may simply dig into the ground to a safe depth (turtle genus Terrapene). In the latter genus, freeze tolerance may play an adaptive role.Most non-marine aquatic turtles overwinter underwater, although Clemmys (Actinemys) marmorata routinely overwinters on land when it occurs in riverine habitats, Kinosternon subrubrum often overwinters on land, and several others may overwinter terrestrially on occasion, especially in more southern climates. For northern species that overwinter underwater, there are two physiological groupings, those that are anoxia-tolerant and those that are relatively anoxia-intolerant. All species fare well physiologically in water with a high partial pressure of oxygen (PO2). A lack of anoxia tolerance limits the types of habitats that a freshwater turtle may live in, since unlike sea turtles, they cannot travel long distances to hibernate.Hatchlings of some species of turtles spend their first winter in or below the nest cavity, while hatchlings of other species in the same area, including northern areas, emerge in the autumn and presumably hibernate underwater. All hatchlings are relatively anoxia-intolerant, and there are no studies to date of where hatchling turtles that do not overwinter in or below the nest cavity spend their first winter. Equally little is known of the ontogeny of anoxia tolerance, other than that adults of all species are more anoxia-tolerant than their hatchlings, probably because of their better ossified shells, which provide adults with more buffer reserves and a larger site in which to sequester lactate. The northern limits of turtles are most likely determined by reproductive limitations (time for egg-laying, incubation, and hatching) than by the rigors of hibernation.Mortality is typically lower in turtle populations during hibernation than it is during their active periods. However, episodic mortality events do occur during hibernation, due to freezing, prolonged anoxia, or predation.     

Effect on airway caliber of stimulation of the hypothalamic locomotor region

Airway dilationis one of the many autonomic responses to exercise. Two neuralmechanisms are believed to evoke these responses: central command andthe muscle reflex. Previously, we found that activation of centralcommand, evoked by electrical and chemical stimulation of themesencephalic locomotor region, constricted the airways rather thandilated them. In the present study we examined in decerebrate paralyzedcats the role played by the hypothalamic locomotor region, theactivation of which also evokes central command, in causing the airwaydilator response to exercise. We found that activation of thehypothalamic locomotor region by electrical and chemical stimuli evokedfictive locomotion and, for the most part, airway constriction. Fictivelocomotion also occurred spontaneously, and this too, for the mostpart, was accompanied by airway constriction. We conclude that centralcommand plays a minor role in the airway dilator response to exercise.

     

Afferent projections to the mesencephalic locomotor region of the cat brain

Afferent projections to the functionally identified mesencephalic locomotor region were investigated in cats using the horseradish peroxidase retrograde axonal transport technique. Sources of afferent projections to this region were discovered in different structures of the fore-, mid-, and hindbrain. Numbers of horseradish peroxidase-labeled neurons were calculated in different brain structures after injecting this enzyme into the mesencephalic locomotor region. Apart from the endopeduncular nucleus, different hypothalamic structures, and the substantia nigra, labeled neurons were discovered in the central tegmental region, the central gray, raphe and vestibular nuclei, the solitary tract nucleus, and the brain stem reticular formation. Neurons accumulating horseradish peroxidase were also discovered in nuclei where ascending sensory tracts originate. This fact serves to bring out the structural inhomogeneity of the midbrain locomotor region; electrical stimulation of this area is an effect which may be attributed to excitation of neurons found within it and activation of accompanying fiber tracts.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev Translated from Neirofiziologiya, Vol. 18, No. 6, pp. 763–773, November–December, 1986.     

Unit responses in the "locomotor strip" of the cat hindbrain to microstimulation

Synaptic responses of single units in the "locomotor strip" of the hindbrain were recorded extracellularly. Short-latency responses appeared in neurons of the rostral part of the strip to stimulation of the "locomotor region" of the mesencephalon. Neurons of the caudal part of the strip responded to microstimulation of its other regions, including rostral. If the distance between the neuron and point of stimulation was under 2–3 mm, short-latency (1.2–1.6 msec) responses could be observed. The thresholds and latent periods of the responses increased when the distance apart increased. Polysynaptic responses with a latent period of 3–4 msec could be potentiated by an increase in the frequency of stimulation up to 30–40 Hz. It is suggested that axons of the "locomotor strip" are oriented in the rostrocaudal direction for a distance of 2–3 mm and give off collaterals which run toward neighboring neurons. The strip may be an integrative center, "intercalated" between the rostral portions of the brain stem and spinal cord.Deceased.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 10, No. 5, pp. 510–518, September–October, 1978.     

Responses of medullary neurons to microstimulation of the "locomotor strip" in cats

Synaptic responses of medullary neurons to stimulation of the bulbar locomotor strip with a current of about 20 µA were studied by an extracellular recording method in mesencephalic cerebellectomized cats. The mean latent period of response of 177 neurons was 3.2 msec. Neurons in which synaptic responses appeared were located in both the lateral and the medial parts of the reticular formation, but short-latency responses were recorded predominantly in the lateral part. In response to a single stimulus 32% of neurons generated a discharge of 2–4 spikes. "Respiratory" neurons were not excited by stimulation of the locomotor point. The results indicate that neurons of the locomotor strip may have an excitatory action not only on each other, but also on neurons located medially. The possible mechanisms of the spread of activity to the superior cervical segments of the spinal cord are discussed.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 13, No. 3, pp. 275–282, May–June, 1981.     

The ontogeny of postmaturation resource allocation in turtles

Resource-allocation decisions vary with life-history strategy, and growing evidence suggests that long-lived endothermic vertebrates direct resources toward growth and self-maintenance when young, increasing allocation toward reproductive effort over time. Few studies have tracked the ontogeny of resource allocation (energy, steroid hormones, etc.) in long-lived ectothermic vertebrates, limiting our understanding of the generality of life-history strategies among vertebrates. We investigated how reproductively mature female painted turtles (Chrysemys picta) from two distinct age classes allocated resources over a 4-yr period and whether resource-allocation patterns varied with nesting experience. We examined age-related variation in body size, egg mass, reproductive frequency, and yolk steroids and report that younger females were smaller and allocated fewer resources to reproduction than did older females. Testosterone levels were higher in eggs from younger females, whereas eggs from second (seasonal) clutches contained higher concentrations of progesterone and estradiol. These allocation patterns resulted in older, larger females laying larger eggs and producing second clutches more frequently than their younger counterparts. We conclude that resource-allocation patterns do vary with age in a long-lived ectotherm.     

Metabolic scaling in turtles

Bennett and Dawson (1976) presented an analysis of the relationship of metabolic rate (MR) and body mass among turtles, based on 10 studies, but unlike most of other groups of ectotherms, there has been no update to include the many later reports on turtles. Here I present a review of the data on turtle metabolic rates at 20, 25, and 30 °C, along with regression equations and graphical analyses from a large number of studies. Two generalities emerge: (1) reported metabolic rates for sea turtles are higher than for other chelonians, although it is not certain whether this is an intrinsic characteristic of sea turtles or an artifact related to experimental conditions (such as greater activity of sea turtles in metabolic chambers and the fact that a number of studies were done with the turtles out of water), and (2) the slopes of the log–log plots of metabolic rate (MR) vs. body mass [b in the allometric equation MR = a(mass)^b] are mostly lower than previously reported in smaller studies.     

Rostral pores in turtles

Rostral pores are epidermal invaginations which occur on the internarial region of most chelonians. Representatives of all Recent chelonian families and 67 of the 74 extant genera were examined grossly and/or histologically. Pores are absent only in the families Carettochelyidae, Cheloniidae, and Dermochelyidae. The number and microscopic structure of pores vary markedly within and between taxa. Morphological data suggest that rostral pores could function in mechanoreception. The possible origin and evolution of rostral pores are discussed in the context of other chelonian integumentary speializations.     

Mitochondrial DNA control region polymorphisms: genetic markers for ecological studies of marine turtles

We describe a rapid and sensitive method for the detection of population-specific genetic markers in mitochondrial DNA (mtDNA) and the use of such markers to analyse population structure of marine turtles. A series of oligonucleotide primers specific for the amplification of the mtDNA control region in Cheloniid turtles were designed from preliminary sequence data. Using two of these primers, a 384–385-bp sequence was amplified from the 5′ portion of the mtDNA control region of 15 green turtles Chelonia mydas from 12 different Indo-Pacific rookeries. Fourteen of the 15 individuals, including some with identical whole-genome restriction fragment patterns, had sequences that differed by one or more base substitutions. Analysis of sequence variation among individuals identified a total of 41 nucleotide substitutions and a 1-bp insertion/deletion. Comparison with evidence from whole-genome restriction enzyme analysis of the same individuals indicated that this portion of the control region is evolving approximately eight times faster than the average rate and that the sequence analysis detected approximately one fifth of the total variation present in the genome. Restriction enzyme analysis of amplified products from an additional 256 individuals revealed significant geographic structuring in the distribution of mtDNA genotypes among five of the 10 rookeries surveyed extensively. Additional geographic structuring of genotypes was identified through denaturing gradient gel electrophoresis (DGGE) of amplified products. Only two of the 10 rookeries surveyed could not be differentiated, indicating that the Indo-Pacific C. mydas include a number of genetically differentiated populations, with minimal female-mediated gene flow among them. Important applications for genetic markers in the conservation and management of marine turtles include the identification of appropriate demographic units for research and management (i.e. genetically discrete populations) and assessment of the composition of feeding and harvested populations.     

Afferent brainstem projections to the hypothalmic locomotor region of the cat brain

The location of sources of direct projections to the hypothalamic locomotor region, electrical stimulation of which in the lightly anesthetized animal induced stepping along a moving treadmill, was studied by the retrograde axonal transport of horseradish peroxidase method in the cat brain stem. Different formations in the brain stem were shown to have direct connections with hypothalamic locomotor regions on both sides. Most sources of these afferent projections were located at sites of catecholamine- (nucl. reticularis lateralis, locus coeruleus, nucl. tractus solitarii) and serotonin-containing (nucl. raphe and substantia grisea centralis) neurons, parabrachial nuclei, and various sensory nuclei. Hypothalamic locomotor regions of both sides form bilateral connections.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 3, pp. 353–362, May–June, 1984.