RESUMEN

The effects of temperature on breathing pattern and oxygen consumption are being investigated in juvenile tortoises and compared to adults, in order to understand physiological adjustments of the respiratory system as related to body size, especially regarding the energetic expenditure associated with growth. We analyzed the breathing pattern and oxygen consumption of juvenile and adult red-footed tortoises (Chelonoidis carbonarius, Cryptodira: Testudinidae). The animals (N = 9; body mass ranging from 0.03 Kg to 2.5 Kg) were exposed to normoxic-normocarbic conditions using open respirometry in order to determine the breathing pattern and oxygen consumption in three different temperatures (15, 25, 35 °C). The obtained results showed intermittent breathing pattern in all tested temperatures in juveniles and adults. Tidal volume was not affected by changes in temperature, while breathing frequency increased significantly with increasing temperature, leading to a significant increment in minute ventilation between 15 and 35 °C. Mass specific oxygen consumption increased significantly with temperature and juveniles showed greater values when compared to adults. The alterations in the ventilatory response to temperature changes occurred in order to maintain the oxygen supply with increased metabolic activity. The differences between juveniles and adults in breathing frequency lead to juveniles needing a lower ventilation rate to perform gas exchange while extracting more oxygen. While these differences might be attributed to a greater metabolic expenditure during development, scaling effects on respiratory variables might be the main contributors to the found differences.

Asunto(s)

Tortugas , Animales , Temperatura , Consumo de Oxígeno/fisiología , Respiración , Oxígeno

RESUMEN

In humans, physical exercise imposes narrower limits for the heart rate (fH) response of the baroreflex, and vascular modulation becomes largely responsible for arterial pressure regulation. In undisturbed reptiles, the baroreflex-related fH alterations at the operating point (Gop) decreases at elevated body temperatures (Tb) and the vascular regulation changes accordingly. We investigated how the baroreflex of rattlesnakes, Crotalus durissus, is regulated during an activity at different Tb, expecting that activity would reduce the capacity of the cardiac baroreflex neural pathway to buffer arterial pressure fluctuations while being compensated by the vascular neural pathway regulation. Snakes were catheterized for blood pressure assessment at three different Tb: 15, 20 and 30 °C. Data were collected before and after activity at each Tb. Baroreflex gain (Gop) was assessed with the sequence method; the vascular limb, with the time constant of pressure decay (τ), using the two-element Windkessel equation. Both Gop and τ reduced when Tb increased. Activity also reduced Gop and τ in all Tb. The relationship between τ and pulse interval (τ/PI) was unaffected by the temperature at resting snakes, albeit it reduced after activity at 20 °C and 30 °C. The unchanged τ/PI and normalized Gop at different Tb indicated those variables are actively adjusted to work at different fH and pressure conditions at rest. Our data suggest that during activity, the baroreflex-related fH response is attenuated and hypertension is buffered by a disproportional increase in the rate which pressure decays during diastole. This compensation seems especially important at higher Tb where Gop is already low.

Asunto(s)

Barorreflejo , Crotalus , Animales , Presión Sanguínea , Frecuencia Cardíaca , Humanos , América del Sur , Temperatura

RESUMEN

Using long-term, remote recordings of heart rate (fH) on fully recovered, undisturbed lizards, we identified several components of heart rate variability (HRV) associated with respiratory sinus arrhythmia (RSA): 1.) A peak in the spectral representation of HRV at the frequency range of ventilation. 2.) These cardiorespiratory interactions were shown to be dependent on the parasympathetic arm of the autonomic nervous system. 3.) Vagal preganglionic neurons are located in discrete groups located in the dorsal motor nucleus of the vagus and also, in a ventro-lateral group, homologous to the nucleus ambiguus of mammals. 4.) Myelinated nerve fibers in the cardiac vagus enabling rapid communication between the central nervous system and the heart. Furthermore, the study of the progressive recovery of fH in tegu following anesthesia and instrumentation revealed that 'resting' levels of mean fH and reestablishment of HRV occurred over different time courses. Accordingly, we suggest that, when an experiment is designed to study a physiological variable reliant on autonomic modulation at its normal, resting level, then postsurgical reestablishment of HRV should be considered as the index of full recovery, rather than mean fH.

Asunto(s)

Sistema Nervioso Autónomo , Frecuencia Cardíaca/fisiología , Corazón/anatomía & histología , Corazón/fisiopatología , Lagartos/fisiología , Recuperación de la Función , Nervio Vago/fisiopatología , Anestesia/métodos , Animales , Masculino , Modelos Teóricos , Respiración , Nervio Vago/anatomía & histología

RESUMEN

The sequence method is an alternative to the traditional pharmacological approach (i.e., the Oxford technique) used to calculate baroreflex gain (G) in mammals. Although the sequence method assesses baroreflex by measuring spontaneous events of blood pressure regulation, the pharmacological method relies on the injection of vasoactive drugs that impact the baroreflex mechanism itself. The sequence method might be relevant for dynamic measurement of baroreflex modulation but it was never validated for vertebrates with low heart rate. Hence, we tested the sequence method in three species of reptiles and compared the results with those provided by the classic pharmacological method. G was similar between both methods and values correlated when parameters for the sequence method were set at delay 0 or 1 (i.e., the baroreflex system responds immediately to blood pressure changes or after 1 heartbeat). Calculation of the baroreflex effectiveness index was adequate at a minimum of 300 cycles and a delay of 1 for the three species. Therefore, the sequence method has been validated to investigate baroreflex regulation in reptiles, enabling studies during dynamic alterations in homeostasis.

Asunto(s)

Barorreflejo/fisiología , Presión Sanguínea/fisiología , Frecuencia Cardíaca/fisiología , Animales , Presión Sanguínea/efectos de los fármacos , Crotalus/fisiología , Iguanas/fisiología , Lagartos/fisiología , Nitroprusiato/farmacología , Fenilefrina/farmacología , Reproducibilidad de los Resultados , Vasoconstrictores/farmacología , Vasodilatadores/farmacología

RESUMEN

This study investigated the maturation of convective oxygen transport in embryos of the snapping turtle (Chelydra serpentina). Measurements included: mass, oxygen consumption (V̇O2 ), heart rate, blood oxygen content and affinity and blood flow distribution at 50%, 70% and 90% of the incubation period. Body mass increased exponentially, paralleled by increased cardiac mass and metabolic rate. Heart rate was constant from 50% to 70% incubation but was significantly reduced at 90% incubation. Hematocrit and hemoglobin concentration were constant at the three points of development studied but arteriovenous difference doubled from 50% to 90% incubation. Oxygen affinity was lower for the early 50% incubation group (stage 19) compared with all other age groups. Blood flow was directed predominantly to the embryo but was highest to the chorioallantoic membrane (CAM) at 70% incubation and was directed away from the yolk as it was depleted at 90% incubation. The findings indicate that the plateau or reduction in egg V̇O2  characteristic of the late incubation period of turtle embryos may be related to an overall reduction in mass-specific V̇O2  that is correlated with decreasing relative heart mass and plateaued CAM blood flow. Importantly, if the blood properties remain unchanged prior to hatching, as they did during the incubation period studied in the current investigation, this could account for the pattern of V̇O2 previously reported for embryonic snapping turtles prior to hatching.

Asunto(s)

Frecuencia Cardíaca , Consumo de Oxígeno , Oxígeno/metabolismo , Transporte Respiratorio , Tortugas/metabolismo , Animales , Peso Corporal , Embrión no Mamífero/metabolismo , Oxígeno/sangre , Tortugas/embriología

RESUMEN

BACKGROUND: Ventilatory parameters have been investigated in several species of Testudines, but few species have had their ventilatory pattern fully characterized by presenting all variables necessary to understand changes in breathing pattern seen under varying environmental conditions. METHODS: We measured ventilation and gas exchange at 25 °C in the semi-aquatic turtle Trachemys scripta and the terrestrial tortoise Chelonoidis carbonarius under normoxia, hypoxia, and hypercarbia and furthermore compiled respiratory data of testudine species from the literature to analyze the relative changes in each variable. RESULTS: During normoxia both species studied showed an episodic breathing pattern with two to three breaths per episode, but the non-ventilatory periods (TNVP) were three to four times longer in T. scripta than in C. carbonarius. Hypoxia and hypercarbia significantly increased ventilation in both species and decreased TNVP and oxygen consumption in T. scripta but not in C. carbonarius. DISCUSSION: Contrary to expectations, the breathing pattern in C. carbonarius did show considerable non-ventilatory periods with more than one breath per breathing episode, and the breathing pattern in T. scripta was found to diverge significantly from predictions based on mechanical analyses of the respiratory system. A quantitative analysis of the literature showed that relative changes in the ventilatory patterns of chelonians in response to hypoxia and hyperbarbia were qualitatively similar among species, although there were variations in the magnitude of change.

RESUMEN

Tegu lizards (Salvator merianae) aestivate for up to 5 months during Brazil's winter, when they retreat to burrows and halt most activities. Dormant tegus reduce their gastrointestinal (GI) mass, which allows a substantial energy economy. This strategy, however, implies that the first post-dormancy digestion would be more costly than subsequent feeding episodes as a result of GI atrophy. To address this, we determined the postprandial metabolic response (SDA) of the first (M1), second (M2) and several (RM) feeding episodes after tegus' dormancy. Another group of tegus (PF) was subjected to an extra 50 day fasting period after arousal. Glucose, triglycerides and uric acid levels were checked before and after feeding. M1 digestion lasted twice as long and cost twofold more when compared with M2 or RM, in agreement with the idea that GI atrophy inflates digestion cost at the first post-dormancy meal. The SDA response was similar in M2 and RM, suggesting that the GI tract was fully reorganized after the first feeding. The SDA cost was equal in PF and RM, implying that the change in state per se (dormant to arousal) triggers the regrowth of GI, independently of feeding. Fasting tegus at M1 presented higher triglyceride and lower uric acid levels than fed tegus, indicating that fasting is mainly sustained by fat storage. Our results show that seasonal fasting imposes an extra digestion cost to tegus following their next feeding, which is fully paid during their first digestion. This surplus cost, however, is negligible compared with the overall energetic savings from GI tract atrophy during the dormancy period.

Asunto(s)

Ayuno/fisiología , Lagartos/metabolismo , Lagartos/fisiología , Animales , Glucemia , Digestión/fisiología , Tracto Gastrointestinal/crecimiento & desarrollo , Tracto Gastrointestinal/fisiología , Hibernación/fisiología , Periodo Posprandial/fisiología , Estaciones del Año , Triglicéridos/sangre , Ácido Úrico/sangre

RESUMEN

The undivided ventricle of non-crocodilian reptiles allows for intracardiac admixture of oxygen-poor and oxygen-rich blood returning via the atria from the systemic circuit and the lungs. The distribution of blood flow between the systemic and pulmonary circuits may vary, based on differences between systemic and pulmonary vascular conductances. The South American rattlesnake, Crotalus durissus, has a single pulmonary artery, innervated by the left vagus. Activity in this nerve controls pulmonary conductance so that left vagotomy abolishes this control. Experimental left vagotomy to abolish cardiac shunting had no effect on long-term survival and failed to identify a functional role in determining metabolic rate, growth or resistance to food deprivation. Accordingly, the present investigation sought to evaluate the extent to which cardiac shunt patterns are actively controlled during changes in body temperature and activity levels. We compared hemodynamic parameters between intact and left-vagotomized rattlesnakes held at different temperatures and subjected to enforced physical activity. Increased temperature and enforced activity raised heart rate, cardiac output, pulmonary and systemic blood flow in both groups, but net cardiac shunt was reversed in the vagotomized group at lower temperatures. We conclude that vagal control of pulmonary conductance is an active mechanism regulating cardiac shunts in C. durissus.

Asunto(s)

Circulación Coronaria/fisiología , Crotalus/fisiología , Nervio Vago/fisiología , Animales , Gasto Cardíaco , Frecuencia Cardíaca , Temperatura , Vagotomía , Nervio Vago/cirugía

RESUMEN

With some notable exceptions, small ectothermic vertebrates are incapable of endogenously sustaining a body temperature substantially above ambient temperature. This view was challenged by our observations of nighttime body temperatures sustained well above ambient (up to 10°C) during the reproductive season in tegu lizards (~2 kg). This led us to hypothesize that tegus have an enhanced capacity to augment heat production and heat conservation. Increased metabolic rates and decreased thermal conductance are the same mechanisms involved in body temperature regulation in those vertebrates traditionally acknowledged as "true endotherms": the birds and mammals. The appreciation that a modern ectotherm the size of the earliest mammals can sustain an elevated body temperature through metabolic rates approaching that of endotherms enlightens the debate over endothermy origins, providing support for the parental care model of endothermy, but not for the assimilation capacity model of endothermy. It also indicates that, contrary to prevailing notions, ectotherms can engage in facultative endothermy, providing a physiological analog in the evolutionary transition to true endothermy.

Asunto(s)

Regulación de la Temperatura Corporal/fisiología , Lagartos/fisiología , Reproducción/fisiología , Termogénesis/fisiología , Adaptación Fisiológica/fisiología , Animales , Aves/fisiología , Temperatura Corporal/fisiología , Metabolismo Energético/fisiología , Mamíferos/fisiología , Estaciones del Año

RESUMEN

Turtles (Testudines) have two major taxa, Pleurodira and Cryptodira. To date, only limited data are available regarding the respiratory physiology of pleurodirans. To begin to address this, we studied ventilation and gas exchange in Podocnemis unifilis and Phrynops geoffroanus. Breathing pattern in both species could be described as episodic with breathing episodes separated by large non-ventilatory periods. We measured duration of inspiration and expiration, breathing frequency, duration of the non-ventilatory period (time between episodes), tidal volume, and oxygen consumption when breathing normoxia, hypoxia and hypercarbia at 25°C. In both species hypercarbia caused a greater increase in ventilation compared to hypoxia, increasing both breathing frequency and tidal volume. Minute ventilation and oxygen consumption in P. geoffroanus were the lowest described so far in testudines, indicating either extra-pulmonary gas exchange or a significantly lower metabolism. Oxidative costs of breathing, estimated using the regression method, was the highest described so far for any reptile. Further studies are necessary to better understand respiratory physiology in Phrynops and Podocnemis species.

Asunto(s)

Fenómenos Fisiológicos Respiratorios , Tortugas/fisiología , Animales , Femenino , Masculino , Especificidad de la Especie

RESUMEN

The autonomic control of heart rate was studied throughout development in embryos of the green iguana, Iguana iguana by applying receptor agonists and antagonists of the parasympathetic and sympathetic systems. Acetylcholine (Ach) slowed or stopped the heart and atropine antagonized the response to Ach indicating the presence of muscarinic cholinoceptors on the heart of early embryos. However, atropine injections had no impact on heart rate until immediately before hatching, when it increased heart rate by 15%. This cholinergic tonus increased to 34% in hatchlings and dropped to 24% in adult iguanas. Although epinephrine was without effect, injection of propranolol slowed the heart throughout development, indicating the presence of ß-adrenergic receptors on the heart of early embryos, possibly stimulated by high levels of circulating catecholamines. The calculated excitatory tonus varied between 33% and 68% until immediately before hatching when it fell to 25% and 29%, a level retained in hatchlings and adults. Hypoxia caused a bradycardia in early embryos that was unaffected by injection of atropine indicating that hypoxia has a direct effect upon the heart. In later embryos and hatchlings hypoxia caused a tachycardia that was unaffected by injection of atropine. Subsequent injection of propranolol reduced heart rate both uncovering a hypoxic bradycardia in late embryos and abolishing tachycardia in hatchlings. Hypercapnia was without effect on heart rate in late stage embryos and in hatchlings.

Asunto(s)

Embrión no Mamífero/fisiología , Frecuencia Cardíaca/fisiología , Corazón/fisiología , Iguanas/fisiología , Acetilcolina/farmacología , Adrenérgicos/farmacología , Agonistas Adrenérgicos beta/farmacología , Antagonistas Adrenérgicos beta/farmacología , Animales , Atropina/farmacología , Sistema Nervioso Autónomo/efectos de los fármacos , Sistema Nervioso Autónomo/embriología , Sistema Nervioso Autónomo/fisiología , Colinérgicos/farmacología , Agonistas Colinérgicos/farmacología , Electrocardiografía , Embrión no Mamífero/efectos de los fármacos , Embrión no Mamífero/embriología , Epinefrina/farmacología , Corazón/efectos de los fármacos , Corazón/embriología , Frecuencia Cardíaca/efectos de los fármacos , Iguanas/embriología , Antagonistas Muscarínicos/farmacología , Miocardio/metabolismo , Propranolol/farmacología , Receptores Adrenérgicos beta/metabolismo , Receptores Colinérgicos/metabolismo

RESUMEN

Measurement of heart rate (fH) in embryonic reptiles has previously imposed some degree of invasive treatment on the developing embryo. Recently a non-invasive technique of fH detection from intact eggs was developed for commercial avian breeders and has since been used in biological research. This device uses infrared light, enabling it to detect heartbeats in very early embryos. However, infrared light is a source of heat and extended enclosure of an egg in the device is likely to affect temperature with consequent effects on physiological processes, including fH. We studied the effect of use of the monitor on the temperature of eggs and on fH in two species of reptiles, the snapping turtle (Chelydra serpentina) and the green iguana (Iguana iguana). Egg temperature increased from a room temperature of 27-28 °C, by 26% in turtles and 14% in iguanas over 1h of enclosure, resulting in an increase in fH of 76-81% in turtles and 35-50% iguanas. These effects on fH can either be avoided by brief enclosure of each egg in the monitor or measured and accounted for during the design of long-term experiments.

Asunto(s)

Técnicas Biosensibles/métodos , Embrión no Mamífero/fisiología , Frecuencia Cardíaca/fisiología , Rayos Infrarrojos , Monitoreo Fisiológico/instrumentación , Animales , Embrión no Mamífero/embriología , Iguanas/embriología , Modelos Lineales , Óvulo/fisiología , Reproducibilidad de los Resultados , Especificidad de la Especie , Temperatura , Factores de Tiempo , Tortugas/embriología

RESUMEN

O crânio representa o segmento com conspícuas adaptações que, nos lagartos, podem ser conservativas ou impulsionadas por pressões seletivas. Objetivando subsidiar o conhecimento morfológico dos répteis, fornecemos uma descrição detalhada dos ossos que formam o neurocrânio de Iguana iguana iguana com base na análise de três esqueletos secos de espécimes adultos. O crânio da referia espécie possui características basais entre os lagartos sem o fechamento das aberturas cranianas e formato geral triangular. As estruturas ósseas que formam a base craniana apresentam muitas fusões, principalmente no assoalho. Na face caudal o exoccipital e o opistótico estão fundidos e formam o otoccipital, que contribui para a formação dos terços laterais do côndilo occipital. A parte central do côndilo é formada pelo supraoccipital. Fusões e estruturas esqueléticas presentes em Iguana são similares aos demais lagartos. Não foram descritas autapomorfias no neurocrânio para esta espécie.(AU)

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Skull represents the segment with conspicuous adaptations that, in lizards, may be conservative or promoted by selective pressures. The aim of assisting the morphological knowledge of reptiles, we provide a detailed description of the neurocranium of Iguana iguana iguana based on analysis of three dried adult skeletons. The skull of this species has basal characteristics in lizards without closure of cranial openings and general triangular shape. Bony structures that form the caudal base have many fusions, especially on the floor. In the caudal face the exoccipital and the opisthotic are fused and form the otooccipital, which contributes to the formation of the lateral part of the condyle. The central part is formed by the condyle supraocciopital. Fusions and skeletal structures in Iguana are similar to other lizards. There are no autopomorphies in the neurocranium for this species.(AU)

Asunto(s)

Animales , Esqueleto/anatomía & histología , Cráneo/anatomía & histología , Iguanas/anatomía & histología

RESUMEN

Heart rate in vertebrates is controlled by activity in the autonomic nervous system. In spontaneously active or experimentally prepared animals, inhibitory parasympathetic control is predominant and is responsible for instantaneous changes in heart rate, such as occur at the first air breath following a period of apnoea in discontinuous breathers like inactive reptiles or species that surface to air breathe after a period of submersion. Parasympathetic control, exerted via fast-conducting, myelinated efferent fibres in the vagus nerve, is also responsible for beat-to-beat changes in heart rate such as the high frequency components observed in spectral analysis of heart rate variability. These include respiratory modulation of the heartbeat that can generate cardiorespiratory synchrony in fish and respiratory sinus arrhythmia in mammals. Both may increase the effectiveness of respiratory gas exchange. Although the central interactions generating respiratory modulation of the heartbeat seem to be highly conserved through vertebrate phylogeny, they are different in kind and location, and in most species are as yet little understood. The heart in vertebrate embryos possesses both muscarinic cholinergic and ß-adrenergic receptors very early in development. Adrenergic control by circulating catecholamines seems important throughout development. However, innervation of the cardiac receptors is delayed and first evidence of a functional cholinergic tonus on the heart, exerted via the vagus nerve, is often seen shortly before or immediately after hatching or birth, suggesting that it may be coordinated with the onset of central respiratory rhythmicity and subsequent breathing.

Asunto(s)

Sistema Nervioso Autónomo/fisiología , Fenómenos Fisiológicos Cardiovasculares , Frecuencia Cardíaca , Filogenia , Fenómenos Fisiológicos Respiratorios , Vertebrados/fisiología , Animales

RESUMEN

Morpho-functional adjustments in the heart of juvenile tegu lizards (Tupinambis merianae) were analyzed at distinct seasonal periods to investigate how the demands of growth and of energy saving are reconciled during the first annual cycle. The relative ventricular mass (Mv) was 31% and 69% larger in late autumn and winter dormancy, respectively, compared to early autumn. This effect did not persist during unfed arousal, suggesting that protein accumulates in the heart during hypometabolism and is degraded on arousal. Both the hypertrophy and the atrophy were disproportionate in the largest individuals. In contrast, Mv was smaller in lizards that were starved during spring activity compared to fed lizards, this effect being larger in smaller individuals. In late autumn and winter dormancy the spongy myocardium had 8% of the section area covered by lacunary spaces, which expanded after food intake during arousal and reached 29% in spring activity together with higher density of cardiomyocytes. Total and soluble proteins per mass unity were unchanged, and maximum activities of selected enzymes suggest sustained glycolytic and aerobic capacities during hypometabolism. Results indicate that important structural adjustments occur in the heart in anticipation of dormancy, and that the protein balance in the tissue is maintained at winter temperatures ~17°C.

Asunto(s)

Cardiomegalia/veterinaria , Lagartos/fisiología , Miocitos Cardíacos/metabolismo , Animales , Índice de Masa Corporal , Cardiomegalia/metabolismo , Cardiomegalia/fisiopatología , Ingestión de Alimentos/fisiología , Metabolismo Energético , Ventrículos Cardíacos/metabolismo , Ventrículos Cardíacos/patología , Lagartos/metabolismo , Miocitos Cardíacos/patología , Estaciones del Año

RESUMEN

The morphologically undivided ventricle of the heart in non-crocodilian reptiles permits the mixing of oxygen-rich blood returning from the lungs and oxygen-poor blood from the systemic circulation. A possible functional significance for this intra-cardiac shunt has been debated for almost a century. Unilateral left vagotomy rendered the single effective pulmonary artery of the South American rattlesnake, Crotalus durissus, unable to adjust the magnitude of blood flow to the lung. The higher constant perfusion of the lung circulation and the incapability of adjusting the right-left shunt in left-denervated snakes persisted over time, providing a unique model for investigation of the long-term consequences of cardiac shunting in a squamate. Oxygen uptake recorded at rest and during spontaneous and forced activity was not affected by removing control of the cardiac shunt. Furthermore, metabolic rate and energetic balance during the post-prandial metabolic increment, plus the food conversion efficiency and growth rate, were all similarly unaffected. These results show that control of cardiac shunting is not associated with a clear functional advantage in adjusting metabolic rate, effectiveness of digestion or growth rates.

Asunto(s)

Crotalus/crecimiento & desarrollo , Crotalus/fisiología , Corazón/fisiología , Consumo de Oxígeno/fisiología , Nervio Vago/fisiología , Anestesia , Animales , Metabolismo Basal/fisiología , Peso Corporal , Estimulación Eléctrica , Conducta Alimentaria/fisiología , Reproducibilidad de los Resultados , Descanso/fisiología , Vagotomía , Nervio Vago/cirugía

RESUMEN

The contribution of air breathing to aerobic metabolic scope and exercise performance was investigated in a teleost with bimodal respiration, the banded knifefish, submitted to a critical swimming speed (U(crit)) protocol at 30°C. Seven individuals (mean ± s.e.m. mass 89±7 g, total length 230±4 mm) achieved a U(crit) of 2.1±1 body lengths (BL) s(-1) and an active metabolic rate (AMR) of 350±21 mg kg(-1) h(-1), with 38±6% derived from air breathing. All of the knifefish exhibited a significant increase in air-breathing frequency (f(AB)) with swimming speed. If denied access to air in normoxia, these individuals achieved a U(crit) of 2.0±0.2 BL s(-1) and an AMR of 368±24 mg kg(-1) h(-1) by gill ventilation alone. In normoxia, therefore, the contribution of air breathing to scope and exercise was entirely facultative. In aquatic hypoxia (P(O(2))=4 kPa) with access to normoxic air, the knifefish achieved a U(crit) of 2.0±0.1 BL s(-1) and an AMR of 338±29 mg kg(-1) h(-1), similar to aquatic normoxia, but with 55±5% of AMR derived from air breathing. Indeed, f(AB) was higher than in normoxia at all swimming speeds, with a profound exponential increase during exercise. If the knifefish were denied access to air in hypoxia, U(crit) declined to 1.2±0.1 BL s(-1) and AMR declined to 199±29 mg kg(-1) h(-1). Therefore, air breathing allowed the knifefish to avoid limitations to aerobic scope and exercise performance in aquatic hypoxia.

Asunto(s)

Aire , Gymnotiformes/fisiología , Condicionamiento Físico Animal , Respiración , Aerobiosis , Animales , Hipoxia , Consumo de Oxígeno , Natación

RESUMEN

Most anatomical and physiological studies of the sauropsid heart have focused on species with extraordinary physiologies, and detailed anatomical descriptions of hearts from sauropsids with more common physiologies are therefore warranted. Here, we present a comprehensive study of the cardiac anatomy of the South American rattlesnake (Crotalus durissus). The cardiovascular physiology of this species has been investigated in a number of studies, whereas only a few cursory studies exist on the cardiac anatomy of viperid snakes. The heart of C. durissus is typically squamate in many regards. Both atria are thin-walled sacs, and the right atrium is the most voluminous. The single ventricle contains three major septa; the vertical septum, the muscular ridge (MR), and the bulbuslamelle. These partially divide the ventricle into three chambers; the systemic and left-sided cavum arteriosum (CA), the pulmonary and right-sided cavum pulmonale, and the medial cavum venosum (CV). The MR is the most developed septum, and several additional and minor septa are found within the CA and CV. An extraordinary thin cortical layer encloses the ventricle, and it is irrigated by a remarkably rich arborization of coronary arteries. Previous studies show high degrees of blood flow separation in the Crotalus heart, and this can only be explained by the coordinated actions of the septa and the prominent atrioventricular valves.

Asunto(s)

Crotalus/anatomía & histología , Corazón/anatomía & histología , Animales , Vasos Coronarios/anatomía & histología , Válvulas Cardíacas/anatomía & histología , América del Sur

RESUMEN

Autonomic control of the cardiovascular system in reptiles includes sympathetic components but heart rate (f(H)), pulmonary blood flow (Q(pul)) and cardiac shunt patterns are primarily controlled by the parasympathetic nervous system. The vagus innervates both the heart and a sphincter on the pulmonary artery. The present study reveals that whereas both the left and right vagi influence f(H), it is only the left vagus that influences pulmonary vascular resistance. This is associated with the fact that rattlesnakes, in common with some other species of snakes, have a single functional lung, as the other lung regresses during development. Stimulation of the left cervical vagus in anaesthetised snakes slowed the heart and markedly reduced blood flow in the pulmonary artery whereas stimulation of the right cervical vagus slowed the heart and caused a small increase in stroke volume (V(S)) in both the systemic and pulmonary circulations. Central stimulation of either vagus caused small (5-10%) reductions in systemic blood pressure but did not affect blood flows or f(H). A bilateral differentiation between the vagi was confirmed by progressive vagotomy in recovered snakes. Transection of the left vagus caused a slight increase in f(H) (10%) but a 70% increase in Q(pul), largely due to an increase in pulmonary stroke volume (V(S,pul)). Subsequent complete vagotomy caused a 60% increase in f(H) accompanied by a slight rise in Q(pul), with no further change in V(S,pul). By contrast, transection of the right vagus elicited a slight tachycardia but no change in V(S,pul). Subsequent complete vagotomy was accompanied by marked increases in f(H), Q(pul) and V(S,pul). These data show that although the heart receives bilateral vagal innervation, the sphincter on the pulmonary artery is innervated solely by the left vagus. This paves the way for an investigation of the role of the cardiac shunt in regulating metabolic rate, as chronic left vagotomy will cause a pronounced left-right shunt in recovered animals, whilst leaving intact control of the heart, via the right vagus.

Asunto(s)

Crotalus/fisiología , Corazón/inervación , Arteria Pulmonar/inervación , Nervio Vago/fisiología , Análisis de Varianza , Animales , Velocidad del Flujo Sanguíneo , Presión Sanguínea , Corazón/fisiología , Frecuencia Cardíaca , Arteria Pulmonar/fisiología , Volumen Sistólico , Vagotomía , Nervio Vago/cirugía

RESUMEN

The reptilian heart consists of a thick inner spongy myocardium that derives its oxygen and nutrient supply directly from the blood within the ventricular cavity, which is surrounded by a thin outer compact layer supplied by coronary arteries. The functional importance of these coronary arteries remains unknown. In the present study we investigate the effects of permanent coronary artery occlusion in the South American rattlesnake (Crotalus durissus) on the ability to maintain heart rate and blood pressure at rest and during short term activity. We used colored silicone rubber (Microfil) to identify the coronary artery distribution and interarterial anastomoses. The coronary circulation was occluded and the snakes were then kept for 4 days at 30 degrees C. Microfil injections verified that virtually all coronary arteries had successfully been occluded, but also made visible an extensive coronary supply to the outer compact layer in untreated snakes. Electrocardiogram (ECG), blood pressure (Psys) and heart rate (fH) were measured at rest and during enforced activity at day 1 and 4. Four days after occlusion of the coronary circulation, the snakes could still maintain a Psys and fH of 5.2+/-0.2 kPa and 58.2+/-2.2 beats min(-1), respectively, during activity and the ECG was not affected. This was not different from sham-operated snakes. Thus, while the outer compact layer of the rattlesnake heart clearly has an extensive coronary supply, rattlesnakes sustain a high blood pressure and heart rate during activity without coronary artery blood supply.

Asunto(s)

Circulación Coronaria , Crotalus/fisiología , Corazón/fisiología , Animales , Presión Sanguínea , Oclusión Coronaria/complicaciones , Vasos Coronarios/anatomía & histología , Vasos Coronarios/fisiología , Crotalus/anatomía & histología , Electrocardiografía , Femenino , Corazón/anatomía & histología , Frecuencia Cardíaca , Hemodinámica , Masculino , América del Sur