The time-dependent effects of hypoxia on the discharge rate carotid chemoreceptors were measured in anesthetized cats. Hypoxic exposure of two different durations were used: a short-term exposure (2-3 h) was used to measure the response of the same carotid chemoreceptors; and a long-term exposure (28 days at inspired PO2 of 70 Torr) to study carotid chemoreceptor properties in one group of cats relative to those of a control group. In the chronically hypoxic and control groups, determinations were made of the 1) steady-state responses to four levels of arterial PO2 (PaO2) at constant levels of arterial PCO2; 2) steady-state responses to acute hypercapnia during hyperoxia; and 3) maximal discharge rates during anoxia. We found that the acute responses of carotid chemoreceptor afferents to a given level of hypoxia (PaO2 = 30-40 Torr) did not significantly change within 2-3 h. After long-term exposure the carotid chemoreceptor responses to hypoxia significantly increased, with no significant changes in the hypercapnic response and in the maximal discharge rate during anoxia. We conclude that isocapnic hypoxia may not elicit a sufficient cellular response within 2-3 h in the cat carotid body to sensitize the O2 responsive mechanism, but hypoxia of longer duration will sensitize such a mechanism, thereby augmenting the chemosensory activity.
Damselfly larvae may autotomize and regenerate any of their 3 caudal lamellae. At least one missing or regenerating lamella was evident in 50.1% of field collected Ischnura posita larvae. Lamellae loss during molting is very infrequent (1 out of 117 recorded molts). Laboratory trials indicate that conspecifics remove lamellae and that this process is density dependent. The percentage of larvae losing lamellae during 24 h trials ranged from 73.5 at the highest density tested to 17.3 at the lowest density. I. posita larvae are cannibalistic. The presence of lamellae reduces an individual's chance of being cannibalized. More than twice as many final instar lamellae-less larvae were cannibalized during 24 h trials than analogous individuals having 3 lamellae at experimental initiation. Costs are also associated with lamellae autotomy. 1) Although individuals without lamellae can swim they are more reluctant to release from a wooden stalk and swim when threatened (9% release) than are larvae with lamellae (29% release). Since swimming is part of their repertoire of anti-predator behaviors this behavioral shift should be detrimental. 2) Caudal lamellae function in O uptake. Trials were conducted with larvae having and not having lamellae in an experimental horizontal oxygen gradient system. Relative to larvae without lamellae, those with lamellae preferred deeper depths at PO values greater than 70 torr. Many lamellae-less larvae distributed themselves at the water surface throughout the range of PO values tested. Differential depth distribution between larvae with and without lamellae is highly significant (P < 0.01).
Interactions between internal and external O2 stimulus levels were assessed by measuring the ventilatory and cardiovascular responses to varying water (PWO2) and air bladder (PabO2) O2 levels and intravascular NaCN in anesthetized spontaneously ventilating Lepisosteus osseus. As PWO2 fell, air-breathing frequency (fab) increased. Buccal pressure amplitude (Pb) also increased as PWO2 fell from hyperoxia to normoxia, but hypoxic water depressed Pb. The PO2 in the ventral aorta (VA) fell as PabO2 fell, which stimulated fab and Pb when the gar was in normoxic or hyperoxic water. Thus gill ventilation and air breathing were normally controlled by both internal and external O2 levels, but aquatic hypoxia uniformly depressed gill ventilation regardless of changes in PabO2 levels. Heart rate and blood pressure were unaffected by these changes. NaCN stimulated hypoxic reflexes and bradycardia more quickly when given into the VA or conus than when given into the dorsal aorta. The animals appear to possess internal chemoreceptors that set the level of hypoxic drive and external chemoreceptors that inhibit gill ventilation and shift the ventilatory emphasis from water to air breathing.
The relative contributions of O2- and CO2-sensitive chemoreceptor information to centrally generated respiratory patterns have changed dramatically during vertebrate evolution. Chemoafferent input from branchial O2 chemoreceptors modulates centrally generated respiratory patterns but is not critical for respiratory rhythmogenesis in fishes. In air-breathing fishes, branchial O2 chemoreceptors monitoring internal and external stimuli control the relative contributions of the gills and air-breathing organ to net ventilation, and chemoafferent input is necessary for initiating air breathing. In the transition from water to air breathing by amphibious vertebrates, rhythmic patterns of branchial ventilation are completely replaced by arrhythmic and intermittent patterns of air breathing, and there is progressive dependence on CO2 as a source of respiratory drive. Periodic initiation of air breathing in resting animals appears to depend on attaining a threshold level of afferent activity from O2- and CO2/pH-sensitive chemoreceptors, since hyperoxia and/or hypocapnia can abolish air breathing in all air-breathing vertebrates. Conversely, chemoreceptor stimulation in amphibians and reptiles converts intermittent to more continuous air breathing patterns, suggesting that adequate biasing input from chemoreceptors activates a central rhythm generator. Chemoafferent input in homeotherms serves as one of several sources of drive for rhythmic breathing and supplies feedback for blood gas homeostasis in the face of metabolic or environmental change.
The transition from water breathing to air breathing for most bimodally breathing fishes appears to be critically dependent on sensory information from three major sets of peripheral receptors. Dominant control over the respiratory mode arises from stimulation of oxygen-sensitive chemoreceptors. Stimulation of internally oriented chemoreceptors generally increases both aquatic and aerial respiration, while stimulation of external chemoreceptors may shift the ventilatory emphasis from water to air breathing. Air-breathing organ mechanoreceptors may help to reflexively stimulate or inhibit air breathing upon deflation or inflation of the air-breathing organ, and probably play a major role in matching ventilation to perfusion in the air-breathing organ. Waterborne irritants or emersion stimulate defense receptors that may override control priorities set by other receptors, and inhibit branchial ventilation in favor of air breathing. While there is still little detailed information about the distribution and characteristics of these sensory receptors, it seems likely that similar sets of receptors control the respiratory mode in most air-breathing fishes, and that differences in the central integration of this sensory information may best account for the great variability of respiratory reflex responses in this diverse group of animals.
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