Evidence for the entrainment of breathing by locomotor pattern in human

In human, it has been shown that interactions between locomotor and respiratory patterns may lead to locomotor-respiratory couplings termed entrainment. In order to prove that this coupling is really an entrainment, we tried to show that it obeys one of the expected rules, i.e. that it evolves and is not present for all imposed locomotor frequencies. For that purpose, seventeen healthy volunteers were asked to run on a treadmill at 14 different locomotor rates (instead of 2 or 3 in previous works) for 40 s. All the subjects did not exhibit the same coupling and different relationships could be obtained: the most commonly observed was 2:1 (2 locomotor activities for a respiratory one) but other forms could appear (4:1 and even 5:2 or 3:2). When the coupling evolution was followed in the same subject, it did not appear for all locomotor frequencies but only for locomotor periods close to harmonics of respiratory ones (absolute coordination). On both sides of these values, it progressively evolved to relative coordination and to the lack of coordination. When two forms of absolute coordination were observed in a same subject, the phase relationships followed the rules of the entrainment. Compared to data obtained in quadrupeds, these results suggest that the entrainment of breathing frequency by the locomotor activity is due to central interactions between the respiratory and locomotor pattern generators and does not depend on a chemical regulation avoided here by short locomotor sequences.     

Cardiac and respiratory activity at arousal from sleep under controlled ventilation conditions.

Arousal from sleep is associated with elevated cardiac and respiratory activity. It is unclear whether this occurs because of homeostatic mechanisms or a reflex activation response associated with arousal. Cardiorespiratory activity was measured during spontaneous arousals from sleep in subjects breathing passively on a ventilator. Under such conditions, homeostatic mechanisms are eliminated. Ventilation, end-tidal PCO2, mask pressure, diaphragmatic electromyograph, heart rate, and blood pressure were measured in four normal subjects under two conditions: assisted ventilation and a normal ventilation control condition. In the control condition, there was a normal, sleep-related fall in ventilation and rise in end-tidal PCO2. Subsequently, at an arousal, there was an increase in respiratory and cardiac activity. In the ventilator condition, a vigorous cardiorespiratory response to a spontaneous arousal from sleep remained. These results indicate that sleep-related respiratory stimuli are not necessary for the occurrence of elevated cardiorespiratory activity at an arousal from sleep and are consistent with the hypothesis that such activity is at least in part due to a reflex activation response.     

Coupling between respiratory and stepping rhythms during locomotion in decerebrate cats

To determine whether and how the strength of coupling between respiratory and stepping rhythms varies depending on locomotor patterns, correlation analysis was done of diaphragmatic and gastrocnemius muscle activities. In spontaneously breathing cats decerebrated at the precollicular-post-mammillary level, tonic electrical stimulation was delivered to the mesencephalic locomotor region to induce locomotion on a treadmill. Electromyograms were recorded from the left hemidiaphragm and the bilateral gastrocnemius muscles. Various locomotor patterns were elicited by changes in the belt speed of the treadmill and in the intensity of stimulation of the mesencephalic locomotor region. Cross-correlograms between diaphragmatic and gastrocnemius activities showed that coupling was absent or weak when the cats walked slowly. The strength of locomotor-respiratory coupling tended to increase as the mean stepping interval shortened. When the animals were galloping, the respiratory rhythm was entrained 1:1 with the stepping rhythm. This study showed that the strength of coupling between respiratory and stepping rhythms varied depending on the locomotor patterns elicited, especially on whether the animals were running.     

Respiratory chemoreceptor function in vertebrates comparative and evolutionary aspects

The sensing of blood gas tensions and/or pH is an evolutionarilyconserved, homeostatic mechanism, observable in almost all speciesstudied from invertebrates to man. In vertebrates, a shift fromthe peripheral O₂-oriented sensing in fish, to the central CO₂/pHsensing in most tetrapods reflects the specific behavioral requirementsof these two groups whereby, in teleost fish, a highly O₂-orientedcontrol of breathing matches the ever-changing and low oxygenlevels in water, whilst the transition to air-breathing increasedthe importance of acid–base regulation and O₂-relateddrive, although retained, became relatively less important.The South American lungfish and tetrapods are probably sistergroups, a conclusion backed up by many similar features of respiratorycontrol. For example, the relative roles of peripheral and centralchemoreceptors are present both in the lungfish and in landvertebrates. In both groups, the central CO₂/pH receptors dominatethe ventilatory response to hypercarbia (60–80%), whilethe peripheral CO₂/pH receptors account for 20–30%. Somebasic components of respiratory control have changed littleduring evolution. This review presents studies that reflectthe current trends in the field of chemoreceptor function, andseveral laboratories are involved. An exhaustive review on theprevious literature, however, is beyond the intended scope ofthe article. Rather, we present examples of current trends inrespiratory function in vertebrates, ranging from fish to humans,and focus on both O₂ sensing and CO₂ sensing. As well, we considerthe impact of chronic levels of hypoxia—a physiologicalcondition in fish and in land vertebrates resident at high elevationsor suffering from one of the many cardiorespiratory diseasestates that predispose an animal to impaired ventilation orcardiac output. This provides a basis for a comparative physiologythat is informative about the evolution of respiratory functionsin vertebrates and about human disease. Currently, most detailis known for mammals, for which molecular biology and respiratoryphysiology have combined in the discovery of the mechanismsunderlying the responses of respiratory chemoreceptors. Ourreview includes new data on nonmammalian vertebrates, whichstresses that some chemoreceptor sites are of ancient origin.     

Breath-by-breath analysis of cardiorespiratory interaction for quantifying developmental maturity in premature infants

In healthy neonates, connections between the heart and lungs through brain stem chemosensory pathways and the autonomic nervous system result in cardiorespiratory synchronization. This interdependence between cardiac and respiratory dynamics can be difficult to measure because of intermittent signal quality in intensive care settings and variability of heart and breathing rates. We employed a phase-based measure suggested by Sch?fer and coworkers (Sch?fer C, Rosenblum MG, Kurths J, Abel HH. Nature 392: 239-240, 1998) to obtain a breath-by-breath analysis of cardiorespiratory interaction. This measure of cardiorespiratory interaction does not distinguish between cardiac control of respiration associated with cardioventilatory coupling and respiratory influences on the heart rate associated with respiratory sinus arrhythmia. We calculated, in sliding 4-min windows, the probability density of heartbeats as a function of the concurrent phase of the respiratory cycle. Probability density functions whose Shannon entropy had a <0.1% chance of occurring from random numbers were classified as exhibiting interaction. In this way, we analyzed 18 infant-years of data from 1,202 patients in the Neonatal Intensive Care Unit at University of Virginia. We found evidence of interaction in 3.3 patient-years of data (18%). Cardiorespiratory interaction increased several-fold with postnatal development, but, surprisingly, the rate of increase was not affected by gestational age at birth. We find evidence for moderate correspondence between this measure of cardiorespiratory interaction and cardioventilatory coupling and no evidence for respiratory sinus arrhythmia, leading to the need for further investigation of the underlying mechanism. Such continuous measures of physiological interaction may serve to gauge developmental maturity in neonatal intensive care patients and prove useful in decisions about incipient illness and about hospital discharge.     

Ecological and Evolutionary Aspects of Integumentary Respiration: Body Size, Diffusion, and the Invertebrata

SYNOPSIS. Most animal phyla lack specialized respiratory surfacesand all phyla contain groups that, for some part of their lifehistory, depend entirely upon integumental diffusion of respiratorygases. Animals that are diffusion-limited, yet function aerobicallyare generally small with large surface areas and there has beenconvergence for this among all phyla including the coelomateinvertebrates. Acoelomates lack specialized respiratory structuresbut have highly modified integuments, functional specializations,and features ranging from symbioses to air gulping that compensatefor diffusion limitation. The diversity of structures functioningfor integumentary respiration is much greater among invertebratesthan vertebrates. Among the higher invertebrates with respiratorysurfaces, accessory integumentary O₂ uptake is usually 20 to50% of total respiration. The high diffusion constant of O₂in air minimizes boundary effects on gas transfer and permitslarger body size, although this is limited by dry conditions.Terrestrial annelids and flatworms, both confined to moist habitats,are larger than aquatic forms which often have accessory gills.Size differences between terrestrial forms in these two phylareflect the presence of a circulation in the annelids. Ontogenetictransitions from skin breathing to other respiratory structuresoccur among marine invertebrates and vertebrates. Vertebratesapparently exercise greater integratory control over integumentalrespiration through adjustment of ventilation and perfusion;however, it is not known if these processes occur in some invertebrates.     

Flexibility in the patterning and control of axial locomotor networks in lamprey

In lower vertebrates, locomotor burst generators for axial muscles generally produce unitary bursts that alternate between the two sides of the body. In lamprey, a lower vertebrate, locomotor activity in the axial ventral roots of the isolated spinal cord can exhibit flexibility in the timings of bursts to dorsally-located myotomal muscle fibers versus ventrally-located myotomal muscle fibers. These episodes of decreased synchrony can occur spontaneously, especially in the rostral spinal cord where the propagating body waves of swimming originate. Application of serotonin, an endogenous spinal neurotransmitter known to presynaptically inhibit excitatory synapses in lamprey, can promote decreased synchrony of dorsal-ventral bursting. These observations suggest the possible existence of dorsal and ventral locomotor networks with modifiable coupling strength between them. Intracellular recordings of motoneurons during locomotor activity provide some support for this model. Pairs of motoneurons innervating myotomal muscle fibers of similar ipsilateral dorsoventral location tend to have higher correlations of fast synaptic activity during fictive locomotion than do pairs of motoneurons innervating myotomes of different ipsilateral dorsoventral locations, suggesting their control by different populations of premotor interneurons. Further, these different motoneuron pools receive different patterns of excitatory and inhibitory inputs from individual reticulospinal neurons, conveyed in part by different sets of premotor interneurons. Perhaps, then, the locomotor network of the lamprey is not simply a unitary burst generator on each side of the spinal cord that activates all ipsilateral body muscles simultaneously. Instead, the burst generator on each side may comprise at least two coupled burst generators, one controlling motoneurons innervating dorsal body muscles and one controlling motoneurons innervating ventral body muscles. The coupling strength between these two ipsilateral burst generators may be modifiable and weakening when greater swimming maneuverability is required. Variable coupling of intrasegmental burst generators in the lamprey may be a precursor to the variable coupling of burst generators observed in the control of locomotion in the joints of limbed vertebrates.     

Respiratory Evolution Facilitated the Origin of Pterosaur Flight and Aerial Gigantism

Pterosaurs, enigmatic extinct Mesozoic reptiles, were the first vertebrates to achieve true flapping flight. Various lines of evidence provide strong support for highly efficient wing design, control, and flight capabilities. However, little is known of the pulmonary system that powered flight in pterosaurs. We investigated the structure and function of the pterosaurian breathing apparatus through a broad scale comparative study of respiratory structure and function in living and extinct archosaurs, using computer-assisted tomographic (CT) scanning of pterosaur and bird skeletal remains, cineradiographic (X-ray film) studies of the skeletal breathing pump in extant birds and alligators, and study of skeletal structure in historic fossil specimens. In this report we present various lines of skeletal evidence that indicate that pterosaurs had a highly effective flow-through respiratory system, capable of sustaining powered flight, predating the appearance of an analogous breathing system in birds by approximately seventy million years. Convergent evolution of gigantism in several Cretaceous pterosaur lineages was made possible through body density reduction by expansion of the pulmonary air sac system throughout the trunk and the distal limb girdle skeleton, highlighting the importance of respiratory adaptations in pterosaur evolution, and the dramatic effect of the release of physical constraints on morphological diversification and evolutionary radiation.     

Oscillations of heart rate and respiration synchronize during poetry recitation

The objective of this study was to investigate the synchronization between low-frequency breathing patterns and respiratory sinus arrhythmia (RSA) of heart rate during guided recitation of poetry, i.e., recitation of hexameter verse from ancient Greek literature performed in a therapeutic setting. Twenty healthy volunteers performed three different types of exercises with respect to a cross-sectional comparison: 1). recitation of hexameter verse, 2). controlled breathing, and 3). spontaneous breathing. Each exercise was divided into three successive measurements: a 15-min baseline measurement (S1), 20 min of exercise, and a 15-min effect measurement (S2). Breathing patterns and RSA were derived from respiratory traces and electrocardiograms, respectively, which were recorded simultaneously using an ambulatory device. The synchronization was then quantified by the index gamma, which has been adopted from the analysis of weakly coupled chaotic oscillators. During recitation of hexameter verse, gamma was high, indicating prominent cardiorespiratory synchronization. The controlled breathing exercise showed cardiorespiratory synchronization to a lesser extent and all resting periods (S1 and S2) had even fewer cardiorespiratory synchronization. During spontaneous breathing, cardiorespiratory synchronization was minimal and hardly observable. The results were largely determined by the extent of a low-frequency component in the breathing oscillations that emerged from the design of hexameter recitation. In conclusion, recitation of hexameter verse exerts a strong influence on RSA by a prominent low-frequency component in the breathing pattern, generating a strong cardiorespiratory synchronization.     

Episodic Breathing in Frogs: Converging Hypotheses on Neural Control of Respiration in Air Breathing Vertebrates

SYNOPSIS. The episodic, or intermittent, breathing of frogsand many ectothermic vertebrates results in important fluctuationsof arterial blood gases. This pattern of breathing differs fromthe rhythmic and continuous alternation of inspiration observedin most homeotherms, which maintain O₂ and CO₂ levels withinnarrow ranges. These differences in pattern of breathing indicatethat the respiratory control systems of ectotherms and homeothermsdiffer substantially. The results of recent studies using invitro brainstemspinal cord preparations of adult frogs and premetamorphictadpoles (Rana catesbeiana and Rana pipiens) demonstrate, however,that the mechanisms for rhythm generation and pattern formationdescribed previously for mammals are also key features of therespiratory control system of frogs. These findings thereforesupport the hypothesis that the respiratory control system ishighly conserved amongst air breathing vertebrates, whetherthey breathe continuously or episodically.     

Medial vestibular nucleus mediates the cardiorespiratory responses to fastigial nuclear activation and hypercapnia.

Electrical stimulation of the cerebellar fastigial nucleus (FN) evokes hyperventilation and hypertension responses that are similar to those induced by stimulation of the medial region of the vestibular nucleus (VNM). Because there are mutual projections between these two nuclei morphologically, we hypothesized that the FN-mediated cardiorespiratory responses were related to the integrity of the VNM. Experiments were conducted on 21 anesthetized, tracheotomized, and spontaneously breathing rats. Electrical stimulation (approximately 10 s) of the FN was used to evoke cardiorespiratory responses, and the same stimulus was repeated 30-45 min after bilateral lesions of the VNM by local microinjection of ibotenic acid (100 mM, 100 nl). We found that FN stimulation-induced hyperventilation and hypertension were attenuated significantly by the lesions. The role of the VNM in the ventilatory responses to chemical challenges was subsequently defined. The animals were exposed to hypercapnia (10% CO2) and hypoxia (10% O2) for 1-2 min randomly before and after VNM lesions. The results showed that VNM lesions significantly attenuated the cardiorespiratory responses to hypercapnia but not to hypoxia, with little effect on baseline respiratory variables. These findings suggest that the VNM is required for full expression of the cardiorespiratory responses to electrical stimulation of the FN as well as to hypercapnia. However, neurons within the VNM do not appear to be critical for maintaining eupneic breathing and the cardiorespiratory responses to hypoxia.     

Amphibians as a model system for the investigation of respiratory control development

Recent medical advances have made it possible for babies to survive premature birth at increasingly earlier developmental stages. This population requires costly and sophisticated medical care to address the problems associated with immaturity of the respiratory system. In addition to pulmonary complications, respiratory instability and apnea reflecting immaturity of the respiratory control system are major causes of hospitalization and morbidity in this highly vulnerable population. These medical concerns, combined with the curiosity of physiologists, have contributed to the expansion of research in respiratory neurobiology. While most researchers working in this field commonly use rodents as an animal model, recent research using in vitro brainstem preparation from bullfrogs (Rana catesbeiana) have revealed the technical advantages of this animal model, and shown that the basic principles underlying respiratory control and its ontogeny are very similar between these two groups of vertebrates. The present review highlights the recent advances in the area of research with a focus on intermittent (episodic) breathing and the role of serotonergic and GABAergic modulation of respiratory activity during development.     

Ontogeny of performance in vertebrates

When competing for food or other resources, or when confronted with predators, young animals may be at a disadvantage relative to adults because of their smaller size. Additionally, the ongoing differentiation and growth of tissues and the development of sensory-motor integration during early ontogeny may constrain performance. Because ectothermic vertebrates show different growth regimes and energetic requirements when compared to endothermic vertebrates, differences in the ontogenetic trajectories of performance traits in these two groups might be expected. However, both groups of vertebrates show similar patterns of changes in performance with ontogeny. Evidence for compensation, resulting in relatively high levels of performance in juveniles relative to adults, appears common for traits related to locomotor and defensive behaviors. However, there is little evidence for compensation in traits associated with feeding and foraging. We suggest that this difference may be due to different selective regimes operating on locomotor versus feeding traits. As a result, relatively high levels of locomotor performance in juveniles and relatively high levels of feeding performance in adults are observed across a wide range of vertebrate groups.     

Respiratory muscle responses elicited by dorsal periaqueductal gray stimulation in rats

The periaqueductal gray matter is an essential neural substrate for central integration of defense behavior and accompanied autonomic responses. The dorsal half of the periaqueductal gray matter (dPAG) is also involved in mediating emotional responses of anxiety and fear, psychological states that often are associated with changes in ventilation. However, information regarding respiratory modulation elicited from this structure is limited. The present study was undertaken to investigate the relationship between stimulus frequency and magnitude on ventilatory pattern and respiratory muscle activity in urethane-anesthetized, spontaneously breathing rats. Electrical stimulation in the dPAG-recruited abdominal muscle activity increased ventilation and increased respiratory frequency by significantly shortening both inspiratory time and expiratory time. Ventilation increased within the first breath after the onset of stimulation, and the respiratory response increased with increasing stimulus frequency and magnitude. dPAG stimulation also increased baseline EMG activity in the diaphragm and recruited baseline external abdominal oblique EMG activity, normally quiescent during eupneic breathing. Significant changes in cardiorespiratory function were only evoked by stimulus intensities >10 microA and when stimulus frequencies were >10 Hz. Respiratory activity of both the diaphragm and abdominal muscles remained elevated for a minimum of 60 s after cessation of stimulation. These results demonstrate that there is a short-latency respiratory response elicited from the dPAG stimulation, which includes both inspiratory and expiratory muscles. The changes in respiratory timing suggest rapid onset and sustained poststimulus dPAG modulation of the brain stem respiratory network that includes expiratory muscle recruitment.     

Inspiring song: The role of respiratory circuitry in the evolution of vertebrate vocal behavior

Vocalization is a common means of communication across vertebrates, but the evolutionary origins of the neural circuits controlling these behaviors are not clear. Peripheral mechanisms of sound production vary widely: fish produce sounds with a swimbladder or pectoral fins; amphibians, reptiles, and mammalians vocalize using a larynx; birds vocalize with a syrinx. Despite the diversity of vocal effectors across taxa, there are many similarities in the neural circuits underlying the control of these organs. Do similarities in vocal circuit structure and function indicate that vocal behaviors first arose in a single common ancestor, or have similar neural circuits arisen independently multiple times during evolution? In this review, we describe the hindbrain circuits that are involved in vocal production across vertebrates. Given that vocalization depends on respiration in most tetrapods, it is not surprising that vocal and respiratory hindbrain circuits across distantly related species are anatomically intermingled and functionally linked. Such vocal‐respiratory circuit integration supports the hypothesis that vocal evolution involved the expansion and functional diversification of breathing circuits. Recent phylogenetic analyses, however, suggest vocal behaviors arose independently in all major tetrapod clades, indicating that similarities in vocal control circuits are the result of repeated co‐options of respiratory circuits in each lineage. It is currently unknown whether vocal circuits across taxa are made up of homologous neurons, or whether vocal neurons in each lineage arose from developmentally and evolutionarily distinct progenitors. Integrative comparative studies of vocal neurons across brain regions and taxa will be required to distinguish between these two scenarios.     

Interseasonal variation in the circadian rhythms of locomotor activity and temperature selection in sleepy lizards, <Emphasis Type="Italic">Tiliqua rugosa</Emphasis>

Few studies in non-mammalian vertebrates have examined how various effectors of the circadian system interact. To determine if the daily locomotor and behavioural thermoregulatory rhythms of Tiliqua rugosa are both controlled by the circadian system in different seasons, lizards were tested in laboratory thermal gradients in four seasons and in constant darkness. Circadian rhythmicity for both rhythms was present in each season, being most pronounced in spring and summer and least evident in autumn. Most lizards displayed a unimodal locomotor activity pattern across all seasons. However, some individuals presented a bimodal locomotor activity pattern in spring and summer. Seasonal variations in the phase relationships of both rhythms to the light:dark (LD) cycle were demonstrated. No seasonal differences in the free-running period lengths of either rhythm were detected, raising the possibility that a single circadian pacemaker drives both rhythms in this species. Our present results demonstrate that both rhythms are similarly controlled by the circadian system in each season. Although seasonal variations in the thermal preferences of reptiles both in the field and laboratory have previously been well documented, this study is the first to demonstrate circadian rhythms of temperature selection in a reptile species in each season.     

Comparative physiology and molecular evolution of carbonic anhydrase in the erythrocytes of early vertebrates

The different isozymes of carbonic anhydrase (CA) have been the subject of intensive study in mammals, but there is still much to be learned about the early evolution of this enzyme in vertebrates. Erythrocyte CA plays an essential role in the respiratory processes of most vertebrates and is probably the most well studied CA isozyme. The available evidence indicates that there has been a progressive increase in the efficiency of erythrocyte CA during the early evolution of vertebrates. There also appears to be a substantial increase in erythrocyte CA activity during development in some species. At the present time, however, the selective pressures that may be influencing the properties of erythrocyte CA during vertebrate evolution and development have not been clearly determined. When the available molecular sequence information is examined, it is evident that the erythrocyte CAs of early vertebrates have active sites that are more similar to those of mammalian CA VII and II, rather than CA I. We can now also begin to examine the phylogenetic relationships between the different rbc CAs in vertebrates, but more CA sequence information is clearly required from different groups of vertebrates before we have a complete picture of the molecular evolution of erythrocyte CA.     

Slow Breathing and Hypoxic Challenge: Cardiorespiratory Consequences and Their Central Neural Substrates

Controlled slow breathing (at 6/min, a rate frequently adopted during yoga practice) can benefit cardiovascular function, including responses to hypoxia. We tested the neural substrates of cardiorespiratory control in humans during volitional controlled breathing and hypoxic challenge using functional magnetic resonance imaging (fMRI). Twenty healthy volunteers were scanned during paced (slow and normal rate) breathing and during spontaneous breathing of normoxic and hypoxic (13% inspired O₂) air. Cardiovascular and respiratory measures were acquired concurrently, including beat-to-beat blood pressure from a subset of participants (N = 7). Slow breathing was associated with increased tidal ventilatory volume. Induced hypoxia raised heart rate and suppressed heart rate variability. Within the brain, slow breathing activated dorsal pons, periaqueductal grey matter, cerebellum, hypothalamus, thalamus and lateral and anterior insular cortices. Blocks of hypoxia activated mid pons, bilateral amygdalae, anterior insular and occipitotemporal cortices. Interaction between slow breathing and hypoxia was expressed in ventral striatal and frontal polar activity. Across conditions, within brainstem, dorsal medullary and pontine activity correlated with tidal volume and inversely with heart rate. Activity in rostroventral medulla correlated with beat-to-beat blood pressure and heart rate variability. Widespread insula and striatal activity tracked decreases in heart rate, while subregions of insular cortex correlated with momentary increases in tidal volume. Our findings define slow breathing effects on central and cardiovascular responses to hypoxic challenge. They highlight the recruitment of discrete brainstem nuclei to cardiorespiratory control, and the engagement of corticostriatal circuitry in support of physiological responses that accompany breathing regulation during hypoxic challenge.     

Cardiorespiratory interactions during resistive load breathing

The addition to the respiratory system of a resistive load results in breathing pattern changes and in negative intrathoracic pressure increases. The aim of this study was to use resistive load breathing as a stimulus to the cardiorespiratory interaction and to examine the extent of the changes in heart rate variability (HRV) and respiratory sinus arrhythmia (RSA) in relation to the breathing pattern changes. HRV and RSA were studied in seven healthy subjects where four resistive loads were applied in a random order during the breath and 8-min recording made in each condition. The HRV spectral power components were computed from the R-R interval sequences, and the RSA amplitude and phase were computed from the sinusoid fitting the instantaneous heart rate within each breath. Adding resistive loads resulted in 1) increasing respiratory period, 2) unchanging heart rate, and 3) increasing HRV and changing RSA characteristics. HRV and RSA characteristics are linearly correlated to the respiratory period. These modifications appear to be linked to load-induced changes in the respiratory period in each individual, because HRV and RSA characteristics are similar at a respiratory period obtained either by loading or by imposed frequency breathing. The present results are discussed with regard to the importance of the breathing cycle duration in these cardiorespiratory interactions, suggesting that these interactions may depend on the time necessary for activation and dissipation of neurotransmitters involved in RSA.     

Properties of Respiratory Pigments in Bimodal Breathing Animals: Air and Water Breathing by Fish and Crustaceans

SYNOPSIS. Oxygen is nearly 30 times more available in air comparedwith water, whereas the carbon dioxide capacity of water is{small tilde}28.5-fold greater than for oxygen, presenting bimodalbreathing species with two very different respiratory milieus.The respiratory pigment plays a variable role in animals switchingbetween the two media. In vertebrates the transition to airbreathing involves two main strategies: a decrease in oxygenaffinity and changes in other haematological parameters suchas haematocrit. When appropriately analyzed, data reveal a decreasein blood oxygen affinity during transition to air. This mayarise via differences in the intrinsic affinity as occurs insome amphibians, or be due to increasesin the organic phosphate:haemoglobin ratio when acclimating to air breathing. Adoptingair breathing often promotes increased haematocrit. It is difficultto discern trends in haemocyanin functioning. Many but not allbimodal and air breathingspecies of crab contain haemocyaninwith high affinity for oxygen. As with haemoglobin there issome tendency for blood haemocyanin concentration to increasewith air breathing but bimodal species are quite variable inthis regard. Different strategies for breathing air are employedby various bimodal crustaceans, some of which involve modulationof haemocyanin oxygenaffinity. The exact mechanisms are oftenspecies dependent and in all bimodal breathing organisms therole of the pigment is best appreciated when the demands ofthe local environment and the behaviour of the species are considered.