RESUMEN
Gill regeneration has not been well studied compared to regeneration of other appendages, such as limb and tail regeneration. Here, we focused on axolotl gill regeneration and found that Fgf- and Bmp-signaling are involved in their gill regeneration mechanism. Axolotls have three pairs of gill rami, and each gill ramus has multiple gill filaments. The gills consist of mesenchyme rich in extracellular matrix and epidermis. The gill nerves are supplied from the trigeminal ganglia located in the head. Denervation resulted in no gill regeneration responses. Nerves and gills express Bmp and Fgf genes, and treating animals with Fgf- and Bmp-signaling inhibitors results in phenotypes similar to those seen in denervated gills. Inducing an accessory appendage is a standard assay in amphibian regeneration research. In our study, an accessory gill could be induced by lateral wounding, suggesting that thin axon fibers and mesenchymal Fgfs and Bmps contributed to the induction of the accessory structure. Such accessory gill induction was inhibited by the denervation. Exogenous Fgf2+Fgf8+Bmp7, which have been determined to function as a regeneration inducer in urodele amphibians, could compensate for the effects denervation has on accessory blastema formation. Our findings suggest that regeneration of appendages in axolotls is regulated by common Fgf- and Bmp-signaling cascades.
Asunto(s)
Ambystoma mexicanum/metabolismo , Ambystoma mexicanum/fisiología , Proteínas Morfogenéticas Óseas/metabolismo , Factores de Crecimiento de Fibroblastos/metabolismo , Branquias/fisiología , Regeneración/fisiología , Transducción de Señal , Ambystoma mexicanum/genética , Animales , Proteínas Morfogenéticas Óseas/genética , Desnervación , Factores de Crecimiento de Fibroblastos/genética , Regulación del Desarrollo de la Expresión Génica , Branquias/inervación , Organogénesis/genética , Ganglio del Trigémino/metabolismoRESUMEN
The baroreflex is one of the most important regulators of cardiovascular homeostasis in vertebrates. It begins with the monitoring of arterial pressure by baroreceptors, which constantly provide the central nervous system with afferent information about the status of this variable. Any change in arterial pressure relative to its normal state triggers autonomic responses, which are characterized by an inversely proportional change in heart rate and systemic vascular resistance and which tend to restore pressure normality. Although the baroreceptors have been located in mammals and other terrestrial vertebrates, their location in fish is still not completely clear and remains quite controversial. Thus, the objective of this study was to locate the baroreceptors in a teleost, the Colossoma macropomum. To do so, the occurrence and efficiency of the baroreflex were both analyzed when this mechanism was induced by pressure imbalancements in intact fish (IN), first-gill-denervated fish (G1), and total-gill-denervated fish (G4). The pressure imbalances were initiated through the administration of the α1-adrenergic agonist phenylephrine (100 µg kg(-1)) and the α1-adrenergic antagonist prazosin (1 mg kg(-1)). The baroreflex responses were then analyzed using an electrocardiogram that allowed for the measurement of the heart rate, the relationship between pre- and post-pharmacological manipulation heart rates, the time required for maximum chronotropic baroreflex response, and total heart rate variability. The results revealed that the barostatic reflex was attenuated in the G1 group and nonexistent in G4 group, findings which indicate that baroreceptors are exclusively located in the gill arches of C. macropomum.
Asunto(s)
Barorreflejo , Peces/fisiología , Branquias/inervación , Branquias/fisiología , Agonistas de Receptores Adrenérgicos alfa 1/farmacología , Antagonistas de Receptores Adrenérgicos alfa 1/farmacología , Animales , Presión Arterial , Desnervación , Electrocardiografía , Femenino , Frecuencia Cardíaca , Masculino , Fenilefrina/farmacología , Prazosina/farmacología , ReflejoRESUMEN
The aim of the present study was to determine the roles that externally versus internally oriented CO(2)/H(+)-sensitive chemoreceptors might play in promoting cardiorespiratory responses to environmental hypercarbia in the air-breathing fish, Hoplerythrinus unitaeniatus (jeju). Fish were exposed to graded hypercarbia (1, 2.5, 5, 10 and 20% CO(2)) and also to graded levels of environmental acidosis (pH approximately 7.0, 6.0, 5.8, 5.6, 5.3 and 4.7) equal to the pH levels of the hypercarbic water to distinguish the relative roles of CO(2) versus H(+). We also injected boluses of CO(2)-equilibrated solutions (5, 10 and 20% CO(2)) and acid solutions equilibrated to the same pH as the CO(2) boluses into the caudal vein (internal) and buccal cavity (external) to distinguish between internal and external stimuli. The putative location of the chemoreceptors was determined by bilateral denervation of branches of cranial nerves IX (glossopharyngeal) and X (vagus) to the gills. The data indicate that the chemoreceptors eliciting bradycardia, hypertension and gill ventilatory responses (increased frequency and amplitude) to hypercarbia are exclusively branchial, externally oriented and respond specifically to changes in CO(2) and not H(+). Those involved in producing the cardiovascular responses appeared to be distributed across all gill arches while those involved in the gill ventilatory responses were located primarily on the first gill arch. Higher levels of aquatic CO(2) depressed gill ventilation and stimulated air breathing. The chemoreceptors involved in producing air breathing in response to hypercarbia also appeared to be branchial, distributed across all gill arches and responded specifically to changes in aquatic CO(2). This would suggest that chemoreceptor groups with different orientations (blood versus water) are involved in eliciting air-breathing responses to hypercarbia in jeju.
Asunto(s)
Aire , Dióxido de Carbono/metabolismo , Células Quimiorreceptoras/metabolismo , Peces/fisiología , Hipercapnia/metabolismo , Reflejo/fisiología , Respiración , Animales , Fenómenos Fisiológicos Cardiovasculares , Células Quimiorreceptoras/citología , Desnervación , Branquias/inervación , Branquias/fisiología , Concentración de Iones de HidrógenoRESUMEN
The localization, distribution and orientation of O(2) chemoreceptors associated with the control of cardio-respiratory responses were investigated in the neotropical, Hoplias lacerdae. Selective denervation of the cranial nerves (IX and X) was combined with chemical stimulation (NaCN) to characterize the gill O(2) chemoreceptors, and the fish were then exposed to gradual hypoxia to examine the extent of each cardio-respiratory response. Changes in heart rate (f(H)) and ventilation amplitude (V(amp)) were allied with chemoreceptors distributed on both internal and external surfaces of all gill arches, while ventilation rate (f) was allied to the O(2) chemoreceptors located only in the internal surface of the first gill arch. H. lacerdae exposed to gradual hypoxia produced a marked bradycardia (45%) and 50% increase in V(amp), but only a relatively small change in f (32%). Thus, the low f(R) response yet high V(amp) were in accord with the characterization of the O(2) chemoreceptors. Comparing these results from H. lacerdae with hypoxia-tolerant species revealed a relationship existent between general oxygenation of the individual species environment, its cardio-respiratory response to hypoxia and the characterization of O(2) chemoreceptors.
Asunto(s)
Adaptación Fisiológica/genética , Fenómenos Fisiológicos Cardiovasculares , Células Quimiorreceptoras/fisiología , Peces/fisiología , Oxígeno/fisiología , Fenómenos Fisiológicos Respiratorios , Animales , Presión Sanguínea/fisiología , Nervios Craneales/cirugía , Desnervación , Branquias/irrigación sanguínea , Branquias/efectos de los fármacos , Branquias/inervación , Frecuencia Cardíaca/fisiología , Especificidad de Órganos , Cianuro de Sodio/farmacología , Especificidad de la Especie , Estrés Fisiológico/fisiologíaRESUMEN
This study examined the location and distribution of O(2) chemoreceptors involved in cardio-respiratory responses to hypoxia in the neotropical teleost, the pacu (Piaractus mesopotamicus). Intact fish and fish experiencing progressive gill denervation by selective transection of cranial nerves IX and X were exposed to gradual hypoxia and submitted to intrabuccal and intravenous injections of NaCN while their heart rate, ventilation rate and ventilation amplitude were measured. The chemoreceptors producing reflex bradycardia were confined to, but distributed along all gill arches, and were sensitive to O(2) levels in the water and the blood. Ventilatory responses to all stimuli, though modified, continued following gill denervation, however, indicating the presence of internally and externally oriented receptors along all gill arches and either in the pseudobranch or at extra-branchial sites. Chemoreceptors located on the first pair of gill arches and innervated by the glossopharyngeal nerve appeared to attenuate the cardiac and respiratory responses to hypoxia. The data indicate that the location and distribution of cardio-respiratory O(2) receptors are not identical to those in tambaqui (Colossoma macropomum) despite their similar habitats and close phylogenetic lineage, although the differences between the two species could reduce to nothing more than the presence or absence of the pseudobranch.
Asunto(s)
Células Quimiorreceptoras/fisiología , Peces/fisiología , Branquias/fisiología , Hipoxia/fisiopatología , Animales , Presión Sanguínea/fisiología , Fenómenos Fisiológicos Cardiovasculares , Branquias/inervación , Frecuencia Cardíaca/fisiología , Oxígeno/metabolismo , Reflejo/fisiología , Fenómenos Fisiológicos Respiratorios , Cianuro de Sodio/farmacología , Especificidad de la EspecieRESUMEN
We examined the cardiorespiratory responses to 6 h of acute hypercarbia (1, 2.5, and 5% CO(2)) in intact and gill-denervated (bilateral denervation of branchial branches of cranial nerves IX and X) tambaqui, Colossoma macropomum. Intact fish exposed to 1 and 2.5% CO(2) increased respiratory frequency ( f(R)) and ventilation amplitude ( V(AMP)) slowly over a 1- to 3-h period. Denervated fish did not show this response, suggesting that tambaqui possess receptors in the gills that will produce excitatory responses to low levels of hypercarbia (1 and 2.5% CO(2)) if the exposure is prolonged. The cardiac response to stimulation of these receptors with this level of CO(2) was a tachycardia and not a bradycardia. During exposure to 5% CO(2), intact fish increased f(R) and V(AMP), and showed a pronounced bradycardia after 1 h. After 2 h, the heart rate ( f(H)) started to increase, but returned to control values after 6 h. In denervated fish, the increase in f(R) was abolished. The slow increase in V(AMP) and the bradycardia were not abolished, suggesting that these changes arose from extra-branchial receptors. Neither intact nor denervated fish developed the swelling of the lower lip or performed aquatic surface respiration, even after 6 h, suggesting that these are unique responses to hypoxia and not hypercarbia.