RESUMEN
To achieve the accuracy and anti-interference of the motion control of the soft robot more effectively, the motion control strategy of the pneumatic soft bionic robot based on the improved Central Pattern Generator (CPG) is proposed. According to the structure and motion characteristics of the robot, a two-layer neural network topology model for the robot is constructed by coupling 22 Hopfield neuron nonlinear oscillators. Then, based on the Adaptive Neuro-Fuzzy Inference System (ANFIS), the membership functions are offline learned and trained to construct the CPG-ANFIS-PID motion control strategy for the robot. Through simulation research on the impact of CPG-ANFIS-PID input parameters on the swimming performance of the robot, it is verified that the control strategy can quickly respond to input parameter changes between different swimming modes, and stably output smooth and continuous dynamic position signals, which has certain advantages. Then, the motion performance of the robot prototype is analyzed experimentally and compared with the simulation results. The results show that the CPG-ANFIS-PID motion control strategy can output coupled waveform signals stably, and control the executing mechanisms of the pneumatic soft bionic robot to achieve biological rhythms motion propulsion waveforms, confirming that the control strategy has accuracy and anti-interference characteristics, and enable the robot have certain maneuverability, flexibility, and environmental adaptability. The significance of this work lies in establishing a CPG-ANFIS-PID control strategy applicable to pneumatic soft bionic robot and proposing a rhythmic motion control method applicable to pneumatic soft bionic robot.
Asunto(s)
Biónica , Redes Neurales de la Computación , Robótica , Robótica/métodos , Robótica/instrumentación , Generadores de Patrones Centrales/fisiología , Lógica Difusa , Simulación por Computador , Movimiento (Física) , Natación/fisiología , AlgoritmosRESUMEN
The appropriate growth of the neurons, accurate organization of their synapses, and successful neurotransmission are indispensable for sensorimotor activities. These processes are highly dynamic and tightly regulated. Extensive genetic, molecular, physiological, and behavioral studies have identified many molecular candidates and investigated their roles in various neuromuscular processes. In this article, we show that Beadex (Bx), the Drosophila LIM only (LMO) protein, is required for motor activities and neuromuscular growth of Drosophila. The larvae bearing Bx7, a null allele of Bx, and the RNAi-mediated neuronal-specific knockdown of Bx show drastically reduced crawling behavior, a diminished synaptic span of the neuromuscular junctions (NMJs) and an increased spontaneous neuronal firing with altered motor patterns in the central pattern generators (CPGs). Microarray studies identified multiple targets of Beadex that are involved in different cellular and molecular pathways, including those associated with the cytoskeleton and mitochondria that could be responsible for the observed neuromuscular defects. With genetic interaction studies, we further show that Highwire (Hiw), a negative regulator of synaptic growth at the NMJs, negatively regulates Bx, as the latter's deficiency was able to rescue the phenotype of the Hiw null mutant, HiwDN. Thus, our data indicate that Beadex functions downstream of Hiw to regulate the larval synaptic growth and physiology.NEW & NOTEWORTHY A novel role for Beadex (Bx) regulates the larval neuromuscular junction (NMJ) structure and function in a tissue-specific manner. Bx is expressed in a subset of Toll-6-expressing neurons and is involved in regulating synaptic span and physiology, possibly through its negative interaction with Highwire (Hiw). The findings of this study provide insights into the molecular mechanisms underlying NMJ development and function and warrant further investigation to understand the role of Bx in these processes fully.
Asunto(s)
Proteínas de Drosophila , Larva , Unión Neuromuscular , Animales , Unión Neuromuscular/fisiología , Unión Neuromuscular/metabolismo , Unión Neuromuscular/crecimiento & desarrollo , Proteínas de Drosophila/metabolismo , Proteínas de Drosophila/genética , Larva/crecimiento & desarrollo , Proteínas con Dominio LIM/metabolismo , Proteínas con Dominio LIM/genética , Drosophila melanogaster/crecimiento & desarrollo , Generadores de Patrones Centrales/fisiología , Generadores de Patrones Centrales/metabolismo , DrosophilaRESUMEN
In response to a suitably aversive skin stimulus, the marine mollusk Tritonia diomedea launches an escape swim followed by several minutes of high-speed crawling. The two escape behaviors are highly dissimilar: whereas the swim is a muscular behavior involving alternating ventral and dorsal whole body flexions, the crawl is a nonrhythmic gliding behavior mediated by the beating of foot cilia. The serotonergic dorsal swim interneurons (DSIs) are members of the swim central pattern generator (CPG) and also strongly drive crawling. Although the swim network is very well understood, the Tritonia crawling network to date comprises only three neurons: the DSIs and pedal neurons 5 and 21 (Pd5 and Pd21). Since Tritonia's swim network has been suggested to have arisen from a preexisting crawling network, we examined the possible role that another swim CPG neuron, C2, may play in crawling. Because of its complete silence in the postswim crawling period, C2 had not previously been considered to play a role in driving crawling. However, semi-intact preparation experiments demonstrated that a brief C2 spike train surprisingly and strongly drives the foot cilia for â¼30 s, something that cannot be explained by its synaptic connections to Pd5 and Pd21. Voltage-sensitive dye (VSD) imaging in the pedal ganglion identified many candidate crawling motor neurons that fire at an elevated rate after the swim and also revealed several pedal neurons that are strongly excited by C2. It is intriguing that unlike the DSIs, which fire tonically after the swim to drive crawling, C2 does so despite its postswim silence.NEW & NOTEWORTHY Tritonia swim central pattern generator (CPG) neuron C2 surprisingly and strongly drives the early phase of postswim crawling despite being silent during this period. In decades of research, C2 had not been suspected of driving crawling because of its complete silence after the swim. Voltage-sensitive dye imaging revealed that the Tritonia crawling motor network may be much larger than previously known and also revealed that many candidate crawling neurons are excited by C2.
Asunto(s)
Generadores de Patrones Centrales , Interneuronas , Natación , Babosas Marinas Tritonia , Animales , Babosas Marinas Tritonia/fisiología , Generadores de Patrones Centrales/fisiología , Natación/fisiología , Interneuronas/fisiología , Potenciales de Acción/fisiología , Neuronas Motoras/fisiología , Reacción de Fuga/fisiologíaRESUMEN
BACKGROUND: The states of the central nervous system (CNS) can be classified into subcritical, critical, and supercritical states that endow the system with information capacity, transmission capabilities, and dynamic range. A further investigation of the relationship between the CNS and the central pattern generators (CPG) is warranted to provide insight into the mechanisms that govern the locomotion system. METHODS: In this study, we established a fractional-order CPG model based on an extended Hindmarsh-Rose model with time delay. A CNS model was further established using a recurrent excitation-inhibition neuronal network. Coupling between these CNS and CPG models was then explored, demonstrating a potential means by which oscillations generated by a neural network respond to periodic stimuli. RESULTS AND CONCLUSIONS: These simulations yielded two key sets of findings. First, frequency sliding was observed when the CPG was sent to the CNS in the subcritical, critical, and supercritical states with different external stimulus and fractional-order index values, indicating that frequency sliding regulates brain function on multiple spatiotemporal scales when the CPG and CNS are coupled together. The main frequency range for these simulations was observed in the gamma band. Second, with increasing external inputs the coherence index for the CNS decreases, demonstrating that strong external inputs introduce neuronal stochasticity. Neural network synchronization is then reduced, triggering irregular neuronal firing. Together these results provide novel insight into the potential mechanisms that may underlie the locomotion system.
Asunto(s)
Encéfalo , Generadores de Patrones Centrales , Modelos Neurológicos , Generadores de Patrones Centrales/fisiología , Encéfalo/fisiología , Humanos , Animales , Redes Neurales de la Computación , Neuronas/fisiología , Simulación por Computador , Red Nerviosa/fisiologíaRESUMEN
Central pattern generators are circuits generating rhythmic movements, such as walking. The majority of existing computational models of these circuits produce antagonistic output where all neurons within a population spike with a broad burst at about the same neuronal phase with respect to network output. However, experimental recordings reveal that many neurons within these circuits fire sparsely, sometimes as rarely as once within a cycle. Here we address the sparse neuronal firing and develop a model to replicate the behavior of individual neurons within rhythm-generating populations to increase biological plausibility and facilitate new insights into the underlying mechanisms of rhythm generation. The developed network architecture is able to produce sparse firing of individual neurons, creating a novel implementation for exploring the contribution of network architecture on rhythmic output. Furthermore, the introduction of sparse firing of individual neurons within the rhythm-generating circuits is one of the factors that allows for a broad neuronal phase representation of firing at the population level. This moves the model toward recent experimental findings of evenly distributed neuronal firing across phases among individual spinal neurons. The network is tested by methodically iterating select parameters to gain an understanding of how connectivity and the interplay of excitation and inhibition influence the output. This knowledge can be applied in future studies to implement a biologically plausible rhythm-generating circuit for testing biological hypotheses.
Asunto(s)
Potenciales de Acción , Generadores de Patrones Centrales , Modelos Neurológicos , Médula Espinal , Potenciales de Acción/fisiología , Generadores de Patrones Centrales/fisiología , Animales , Médula Espinal/fisiología , Neuronas/fisiología , Simulación por Computador , Redes Neurales de la Computación , Periodicidad , Red Nerviosa/fisiología , HumanosRESUMEN
Swallowing is induced by a central pattern generator in the nucleus tractus solitarius (NTS). We aimed to create a medullary slice preparation to elucidate the neural architecture of the central pattern generator of swallowing (Sw-CPG) and record its neural activities. Experiments were conducted on 2-day-old Sprague-Dawley rats (n = 46). The brainstem-spinal cord was transected at the pontomedullary and cervicothoracic junctions; the medulla was sliced transversely at thicknesses of 600, 700, or 800 µm. The rostral end of the slice was 100 µm rostral to the vagus nerve. We recorded hypoglossal nerve activity and electrically stimulated the vagus nerve or microinjected bicuculline methiodide (BIC) into the NTS. The 800-µm slices generated both rhythmic respiratory activity and electrically elicited neural activity. The 700-µm slices generated only respiratory activity, while the 600-µm slices did not generate any neural activity. BIC microinjection into the NTS in 800-µm slices resulted in the typical activity that closely resembled the swallowing activity reported in other experiments. This swallowing-like activity consistently lengthened the respiratory interval. Despite complete inhibition of respiratory activity, weak swallowing-like activity was observed under bath application of a non-NMDA receptor antagonist. Contrastingly, bath application of NMDA receptor antagonists resulted in a complete loss of swallowing-like activity and no change in respiratory activity. These results suggest that the 800-µm medullary slice preparation contains both afferent and efferent neural circuits and pattern generators of swallowing activity. Additionally, NMDA receptors may be necessary for generating swallowing activity. This medullary slice preparation can therefore elucidate Sw-CPG neural networks.
Asunto(s)
Animales Recién Nacidos , Bicuculina , Generadores de Patrones Centrales , Deglución , Nervio Hipogloso , Bulbo Raquídeo , Ratas Sprague-Dawley , Nervio Vago , Animales , Deglución/fisiología , Deglución/efectos de los fármacos , Bulbo Raquídeo/fisiología , Bulbo Raquídeo/efectos de los fármacos , Bicuculina/farmacología , Bicuculina/análogos & derivados , Ratas , Nervio Vago/fisiología , Nervio Vago/efectos de los fármacos , Generadores de Patrones Centrales/fisiología , Generadores de Patrones Centrales/efectos de los fármacos , Nervio Hipogloso/fisiología , Nervio Hipogloso/efectos de los fármacos , Estimulación Eléctrica , Núcleo Solitario/efectos de los fármacos , Núcleo Solitario/fisiologíaRESUMEN
Central pattern generators are small networks that contribute to generating animal locomotion. The models used to study gait generation and gait transition mechanisms often require biologically accurate neuron and synapse models, with high dimensionality and complex dynamics. Tuning the parameters of these models to elicit network dynamics compatible with gait features is not a trivial task, due to the impossibility of inferring a priori the effects of each parameter on the nonlinear system's emergent dynamics. In this paper we explore the use of global optimization strategies for parameter optimization in multigait central pattern generator (CPG) models with complex cell dynamics and minimal topology. We first consider an existing quadruped CPG model as a test bed for the objective function formulation, then proceed to optimize the parameters of a newly proposed multigait, interlimb hexapod CPG model. We successfully obtain hexapod gaits and prompt gait transitions by varying only control currents, while all CPG parameters, once optimized, are kept fixed. This mechanism of gait transitions is compatible with short-term synaptic plasticity.
Asunto(s)
Generadores de Patrones Centrales , Animales , Generadores de Patrones Centrales/fisiología , Marcha/fisiología , Locomoción/fisiología , Dinámicas no Lineales , NeuronasRESUMEN
Within vertebrates, central pattern generators drive rhythmical behaviours, such as locomotion and ventilation. Their pattern generation is also influenced by sensory input and various forms of neuromodulation. These capabilities arose early in vertebrate evolution, preceding the evolution of the cerebellum in jawed vertebrates. This later evolution of the cerebellum is suggestive of subsumption architecture that adds functionality to a pre-existing network. From a central-pattern-generator perspective, what additional functionality might the cerebellum provide? The suggestion is that the adaptive filter capabilities of the cerebellum may be able to use error learning to appropriately repurpose pattern output. Examples may include head and eye stabilization during locomotion, song learning, and context-dependent alternation between learnt motor-control sequences.
Asunto(s)
Generadores de Patrones Centrales , Cerebelo , Animales , Cerebelo/fisiología , Locomoción/fisiología , Aprendizaje/fisiología , Generadores de Patrones Centrales/fisiologíaRESUMEN
Objective.Studying the neural components regulating movement in human locomotion is obstructed by the inability to perform invasive experimental recording in the human neural circuits. Neuromechanical simulations can provide insights by modeling the locomotor circuits. Past neuromechanical models proposed control of locomotion either driven by central pattern generators (CPGs) with simple sensory commands or by a purely reflex-based network regulated by state-machine mechanisms, which activate and deactivate reflexes depending on the detected gait cycle phases. However, the physiological interpretation of these state machines remains unclear. Here, we present a physiologically plausible model to investigate spinal control and modulation of human locomotion.Approach.We propose a bio-inspired controller composed of two coupled CPGs that produce the rhythm and pattern, and a reflex-based network simulating low-level reflex pathways and Renshaw cells. This reflex network is based on leaky-integration neurons, and the whole system does not rely on changing reflex gains according to the gait cycle state. The musculoskeletal model is composed of a skeletal structure and nine muscles per leg generating movement in sagittal plane.Main results.Optimizing the open parameters for effort minimization and stability, human kinematics and muscle activation naturally emerged. Furthermore, when CPGs were not activated, periodic motion could not be achieved through optimization, suggesting the necessity of this component to generate rhythmic behavior without a state machine mechanism regulating reflex activation. The controller could reproduce ranges of speeds from 0.3 to 1.9 m s-1. The results showed that the net influence of feedback on motoneurons (MNs) during perturbed locomotion is predominantly inhibitory and that the CPGs provide the timing of MNs' activation by exciting or inhibiting muscles in specific gait phases.Significance.The proposed bio-inspired controller could contribute to our understanding of locomotor circuits of the intact spinal cord and could be used to study neuromotor disorders.
Asunto(s)
Generadores de Patrones Centrales , Humanos , Generadores de Patrones Centrales/fisiología , Locomoción/fisiología , Marcha/fisiología , Médula Espinal/fisiología , Reflejo/fisiologíaRESUMEN
The ability to generate and control locomotor movements depends on complex interactions between many areas of the nervous system, the musculoskeletal system, and the environment. How the nervous system manages to accomplish this task has been the subject of investigation for more than a century. In vertebrates, locomotion is generated by neural networks located in the spinal cord referred to as central pattern generators. Descending inputs from the brain stem initiate, maintain, and stop locomotion as well as control speed and direction. Sensory inputs adapt locomotor programs to the environmental conditions. This review presents a comparative and historical overview of some of the neural mechanisms underlying the control of locomotion in vertebrates. We have put an emphasis on spinal mechanisms and descending control.
Asunto(s)
Generadores de Patrones Centrales , Médula Espinal , Animales , Médula Espinal/fisiología , Tronco Encefálico/fisiología , Locomoción/fisiología , Lampreas/fisiología , Redes Neurales de la Computación , Generadores de Patrones Centrales/fisiologíaRESUMEN
Lumbar central pattern generators (CPGs) control the basic rhythm and coordinate muscle activation underlying hindlimb locomotion in quadrupedal mammals. The existence and function of CPGs in humans have remained controversial. Here, we investigated a case of a male individual with complete thoracic spinal cord injury who presented with a rare form of self-sustained rhythmic spinal myoclonus in the legs and rhythmic activities induced by epidural electrical stimulation (EES). Analysis of muscle activation patterns suggested that the myoclonus tapped into spinal circuits that generate muscle spasms, rather than reflecting locomotor CPG activity as previously thought. The EES-induced patterns were fundamentally different in that they included flexor-extensor and left-right alternations, hallmarks of locomotor CPGs, and showed spontaneous errors in rhythmicity. These motor deletions, with preserved cycle frequency and period when rhythmic activity resumed, were previously reported only in animal studies and suggest a separation between rhythm generation and pattern formation. Spinal myoclonus and the EES-induced activity demonstrate that the human lumbar spinal cord contains distinct mechanisms for generating rhythmic multi-muscle patterns.
Asunto(s)
Generadores de Patrones Centrales , Mioclonía , Traumatismos de la Médula Espinal , Animales , Masculino , Humanos , Médula Espinal/fisiología , Locomoción/fisiología , Miembro Posterior , Generadores de Patrones Centrales/fisiología , MamíferosRESUMEN
Across phyla, males often produce species-specific vocalizations to attract females. Although understanding the neural mechanisms underlying behavior has been challenging in vertebrates, we previously identified two anatomically distinct central pattern generators (CPGs) that drive the fast and slow clicks of male Xenopus laevis, using an ex vivo preparation that produces fictive vocalizations. Here, we extended this approach to four additional species, X. amieti, X. cliivi, X. petersii, and X. tropicalis, by developing ex vivo brain preparation from which fictive vocalizations are elicited in response to a chemical or electrical stimulus. We found that even though the courtship calls are species-specific, the CPGs used to generate clicks are conserved across species. The fast CPGs, which critically rely on reciprocal connections between the parabrachial nucleus and the nucleus ambiguus, are conserved among fast-click species, and slow CPGs are shared among slow-click species. In addition, our results suggest that testosterone plays a role in organizing fast CPGs in fast-click species, but not in slow-click species. Moreover, fast CPGs are not inherited by all species but monopolized by fast-click species. The results suggest that species-specific calls of the genus Xenopus have evolved by utilizing conserved slow and/or fast CPGs inherited by each species.
Asunto(s)
Generadores de Patrones Centrales , Animales , Femenino , Masculino , Xenopus laevis/fisiología , Generadores de Patrones Centrales/fisiología , Vocalización Animal/fisiología , Bulbo Raquídeo , Encéfalo/fisiologíaRESUMEN
In the past two decades we have learned an enormous amount of information regarding the identity of functional components of the neural circuitry responsible for generating locomotor activity in mammals. Molecular techniques, combined with classic electrophysiological and anatomical approaches, have resulted in the identification of a handful of classes of genetically defined interneuronal populations, and a delineation of the specific function of many of these during stepping. What lags behind at this point is a clear picture of the synaptic connectivity of each population, this information is key if we are to understand how the interneuronal components that are responsible for locomotor activity work together to form a functional circuit. In this mini review I will summarize what is, and what is not, known regarding the synaptic connectivity of each genetically defined interneuronal population that is involved in locomotion.
Asunto(s)
Generadores de Patrones Centrales , Animales , Generadores de Patrones Centrales/fisiología , Médula Espinal/fisiología , Locomoción/fisiología , Interneuronas/fisiología , Fenómenos Electrofisiológicos , MamíferosRESUMEN
The neurophysiological background of oriental bodywork based on breathing methods is poorly understood. Recently, experimental techniques using genetically engineered animals have been applied to identify and analyze the functions of motor columns such as medial motor column (MMC), known as evolutionarily old locomotion central pattern generators (CPGs) in lamprey. In oriental bodywork between two individuals, locomotion-related body reactions are often observed. In Nishino Breathing Methods (NBM), the extrapyramidal system is likely to be accessed through the practice of Sokushin-kokyu (whole body awareness with breathing), Karin (body axis rotation and reactivation with breathing), and Taiki (body trunk mutual signaling with breathing; the coordination of two individual body trunks). In addition, kinesiological discussions of ancient oriental traditions, such as the practice of mysterious hand-back contact and the active expiration that relates to music and body trunk reaction, are presented. Based on the hypothesis that oriental bodywork is accessing the extrapyramidal body trunk motor system (basal ganglia to spiral locomotion CPGs), this paper discusses the inter-individual body reactions coexisting through different evolutionary levels in multilayered motor systems, and their possible medical applications.
Asunto(s)
Generadores de Patrones Centrales , Manipulaciones Musculoesqueléticas , Animales , Generadores de Patrones Centrales/fisiología , Marcha , Mano , Locomoción/fisiologíaRESUMEN
This chapter reviews recent work showing that vertebrate motoneurons can trigger spontaneous rhythmic activity in the developing spinal cord and can modulate the function of several different central pattern generators later in development. In both the embryonic chick and the fetal mouse spinal cords, antidromic activation of motoneurons can trigger bouts of rhythmic activity. In the neonatal mouse, optogenetic manipulation of motoneuron firing can modulate the frequency of fictive locomotion activated by a drug cocktail. In adult animals, motoneurons have been shown to regulate swimming in the zebrafish, and vocalization in fish and frogs. We discuss the significance of these findings and the degree to which motoneurons may be considered a part of these central pattern generators.
Asunto(s)
Generadores de Patrones Centrales , Animales , Generadores de Patrones Centrales/fisiología , Locomoción/fisiología , Ratones , Neuronas Motoras , Médula Espinal , Pez CebraRESUMEN
Central pattern generators (CPGs) generate the rhythmic and coordinated neural features necessary for the proper conduction of complex behaviors. In particular, CPGs are crucial for complex motor behaviors such as locomotion, mastication, respiration, and vocal production. While the importance of these networks in modulating behavior is evident, the mechanisms driving these CPGs are still not fully understood. On the other hand, accumulating evidence suggests that astrocytes have a significant role in regulating the function of some of these CPGs. Here, we review the location, function, and role of astrocytes in locomotion, respiration, and mastication CPGs and propose that, similarly, astrocytes may also play a significant role in the vocalization CPG.
Asunto(s)
Astrocitos , Generadores de Patrones Centrales , Generadores de Patrones Centrales/fisiología , Locomoción/fisiología , RespiraciónRESUMEN
Astrocytes are thought to play a crucial role in providing structure to the spinal cord and maintaining efficient synaptic function and metabolism because their fine processes envelop the synapses of neurons and form many neuronal networks within the central nervous system (CNS). To investigate whether putative astrocytes and putative neurons distributed on the ventral horn play a role in the modulation of lumbar locomotor central pattern generator (CPG) networks, we used extracellular recording and optical imaging techniques and recorded the neural output from the left L5 ventral root and the calcium activity of putative astrocytes and neurons in the L5 ventral horn at the same time when activating an isolated L1-L5 spinal cord preparation from rats aged 0-2 days. Optical measurements detected cells that showed a fluorescence intensity change under all experimental conditions, namely, (1) 5-HT + NMDA, (2) TTX, and (3) TTX + Low K+. These cells were semiautomatically identified using an in-house MATLAB-based program, as putative astrocytes and neurons according to the cell classification, i.e., increased or decreased fluorescence intensity change (ΔF/F0), and subjective judgment based on their soma size. Coherence and its phase were calculated according to the calcium activity of the putative astrocytes and putative neurons, and neural output was calculated during fictive locomotion with in-house MATLAB-based programs. We found that the number of putative astrocytes activated by applying low K+ tends not to differ from that activated by applying the protease-activated receptor 1 (PAR1) selective agonist TFLLR-NH2 (TFLLR). Moreover, the calcium activity of several putative astrocytes and neurons synchronized with locomotor-like activity at a frequency range below 0.5 Hz and the time lag between peaks of cellular calcium activity and locomotor-like activity ranged from -1000 to + 1000 ms. These findings presumably indicates that these putative astrocytes and neurons in the left L5 ventral horn require -1000 to + 1000 ms to communicate with lumbar CPG networks and maintain efficient synaptic function and metabolism in activated lumbar CPG networks. This finding suggests the possibility that putative astrocytic and neuronal cells in the L5 ventral horn contribute to generating the rhythms and patterns of locomotor-like activity by activated CPG networks in the first to fifth lumbar spinal cord.
Asunto(s)
Células del Asta Anterior/metabolismo , Astrocitos/metabolismo , Señalización del Calcio , Generadores de Patrones Centrales/metabolismo , Locomoción , Animales , Células del Asta Anterior/efectos de los fármacos , Células del Asta Anterior/fisiología , Astrocitos/efectos de los fármacos , Astrocitos/fisiología , Generadores de Patrones Centrales/efectos de los fármacos , Generadores de Patrones Centrales/fisiología , N-Metilaspartato/metabolismo , Oligopéptidos/farmacología , Potasio/metabolismo , Ratas , Ratas Wistar , Serotonina/metabolismo , Tetrodotoxina/farmacologíaRESUMEN
Tracing the evolution of the central rhythm generators associated with ventilation in vertebrates is hindered by a lack of information surrounding key transitions. To begin with, central rhythm generation has been studied in detail in only a few species from four vertebrate groups, lamprey, anuran amphibians, turtles, and mammals (primarily rodents). Secondly, there is a lack of information regarding the transition from water breathing fish to air breathing amniotes (reptiles, birds, and mammals). Specifically, the respiratory rhythm generators of fish appear to be single oscillators capable of generating both phases of the respiratory cycle (expansion and compression) and projecting to motoneurons in cranial nerves innervating bucco-pharyngeal muscles. In the amniotes we find oscillators capable of independently generating separate phases of the respiratory cycle (expiration and inspiration) and projecting to pre-motoneurons in the ventrolateral medulla that in turn project to spinal motoneurons innervating thoracic and abdominal muscles (reptiles, birds, and mammals). Studies of the one group of amphibians that lie at this transition (the anurans), raise intriguing possibilities but, for a variety of reasons that we explore, also raise unanswered questions. In this review we summarize what is known about the rhythm generating circuits associated with breathing that arise from the different rhombomeric segments in each of the different vertebrate classes. Assuming oscillating circuits form in every pair of rhombomeres in every vertebrate during development, we trace what appears to be the evolutionary fate of each and highlight the questions that remain to be answered to properly understand the evolutionary transitions in vertebrate central respiratory rhythm generation.
Asunto(s)
Evolución Biológica , Generadores de Patrones Centrales/fisiología , Respiración , Animales , VertebradosRESUMEN
Central pattern generators (CPGs) are neurons or neural circuits that produce periodic output without requiring patterned input. More complex behaviors can be assembled from simpler subroutines, and nested CPGs have been proposed to coordinate their repetitive elements, organizing control over different time scales. Here, we use behavioral experiments to establish that Drosophila grooming may be controlled by nested CPGs. On a short time scale (5-7 Hz, ~ 200 ms/movement), flies clean with periodic leg sweeps and rubs. More surprisingly, transitions between bouts of head sweeping and leg rubbing are also periodic on a longer time scale (0.3-0.6 Hz, ~2 s/bout). We examine grooming at a range of temperatures to show that the frequencies of both oscillations increase-a hallmark of CPG control-and also that rhythms at the two time scales increase at the same rate, indicating that the nested CPGs may be linked. This relationship holds when sensory drive is held constant using optogenetic activation, but oscillations can decouple in spontaneously grooming flies, showing that alternative control modes are possible. Loss of sensory feedback does not disrupt periodicity but slow down the longer time scale alternation. Nested CPGs simplify the generation of complex but repetitive behaviors, and identifying them in Drosophila grooming presents an opportunity to map the neural circuits that constitute them.
Asunto(s)
Drosophila melanogaster/fisiología , Aseo Animal/fisiología , Movimiento , Animales , Generadores de Patrones Centrales/fisiología , Optogenética , PeriodicidadRESUMEN
Mutually inhibitory populations of neurons, half-center oscillators (HCOs), are commonly involved in the dynamics of the central pattern generators (CPGs) driving various rhythmic movements. Previously, we developed a multifunctional, multistable symmetric HCO model which produced slow locomotor-like and fast paw-shake-like activity patterns. Here, we describe asymmetric features of paw-shake responses in a symmetric HCO model and test these predictions experimentally. We considered bursting properties of the two model half-centers during transient paw-shake-like responses to short perturbations during locomotor-like activity. We found that when a current pulse was applied during the spiking phase of one half-center, let's call it #1, the consecutive burst durations (BDs) of that half-center increased throughout the paw-shake response, while BDs of the other half-center, let's call it #2, only changed slightly. In contrast, the consecutive interburst intervals (IBIs) of half-center #1 changed little, while IBIs of half-center #2 increased. We demonstrated that this asymmetry between the half-centers depends on the phase of the locomotor-like rhythm at which the perturbation was applied. We suggest that the fast transient response reflects functional asymmetries of slow processes that underly the locomotor-like pattern; e.g., asymmetric levels of inactivation across the two half-centers for a slowly inactivating inward current. We compared model results with those of in-vivo paw-shake responses evoked in locomoting cats and found similar asymmetries. Electromyographic (EMG) BDs of anterior hindlimb muscles with flexor-related activity increased in consecutive paw-shake cycles, while BD of posterior muscles with extensor-related activity did not change, and vice versa for IBIs of anterior flexors and posterior extensors. We conclude that EMG activity patterns during paw-shaking are consistent with the proposed mechanism producing transient paw-shake-like bursting patterns found in our multistable HCO model. We suggest that the described asymmetry of paw-shaking responses could implicate a multifunctional CPG controlling both locomotion and paw-shaking.