Dona F. Boggs

Activity of three muscles associated with the uncinate processes of the giant Canada goose Branta canadensis maximus

SUMMARY The activity of three muscles associated with the uncinate processes, (i) external intercostal, (ii) appendicocostalis and (iii) external oblique, were examined using patch and sew-through electrodes during sitting, standing and moderate speed treadmill running in the giant Canada goose. The external intercostal muscles demonstrated no respiratory activity, being active only during running, suggesting they play some role in trunk stabilisation. The appendicocostalis and external oblique muscles are respiratory muscles, being active during inspiration and expiration, respectively. The activity of the appendicocostalis muscles increased during sitting, suggesting that the uncinate processes in birds play an important role in facilitating lateral flaring of the rib cage when sternal rocking is prevented. We suggest that the uncinate processes in birds facilitate movements of the ribs and sternum during breathing and therefore are integral to the breathing mechanics of birds.

Role of theRete mirabile ophthalmicum in maintaining the body-to-brain temperature difference in pigeons

Summary1.Brain (hypothalamic) and colonic temperatures were measured in twenty adult pigeons (Columba livia) whose mean body mass was 0.377 kg.2.In contrals, in sham operated birds, and in those pigeons in which one or both external ophthalmic arteries were occluded brain temperatures were always about 1°C (0.94 to 1.03) below body temperature (Fig. 2) over a range of air temperatures.3.In pigeons in which arterial flow to theretia was totally blocked, the normal pattern of body-to-brain temperature difference wasreversed, such that brain temperature was always higher than body temperature by a mean of 0.36 °C (Fig. 2, Table 1).4.Therete mirabile ophthalmicum of pigeons plays a central role in the maintenance of the body-to-brain temperature difference which may be important in avoiding brain damage during core hyperthermia.

Interactions between locomotion and ventilation in tetrapods

Interactions between locomotion and ventilation have now been studied in several species of reptiles, birds and mammals, from a variety of perspectives. Among these perspectives are neural interactions of separate but linked central controllers; mechanical impacts of locomotion upon ventilatory pressures and flows; and the extent to which the latter may affect gas exchange and the energetics of exercise. A synchrony, i.e. 1:1 pattern of coordination, is observed in many running mammals once they achieve galloping speeds, as well as in flying bats, some flying birds and hopping marsupials. Other, non-1:1, patterns of coordination are seen in trotting and walking quadrupeds, as well as running bipedal humans and running and flying birds. There is evidence for an energetic advantage to coordination of locomotor and respiratory cycles for flying birds and running mammals. There is evidence for a mechanical constraint upon ventilation by locomotion for some reptiles (e.g. iguana), but not for others (e.g. varanids and crocodilians). In diving birds the impact of wing flapping or foot paddling on differential air sac pressures enhances gas exchange during the breath hold by improving diffusive and convective movement of air sac oxygen to parabronchi. This paper will review the current state of our knowledge of such influences of locomotion upon respiratory system function.

Effect of perinatal hypercapnia on the adult ventillatory response to carbon dioxide

Burrowing mammals show a reduced ventilatory response to CO2 and CO2 retention. We examined whether this reduced responsiveness could be due to modification of chemoreceptors by persistent hypercapnia during development. Mice and rats were exposed to 6.0% CO2 throughout gestation and/or weaning and then removed to normocapnic air for a minimum of 6 weeks. Mouse gas pocket O2 and CO2 tensions and hematocrits were analyzed and compared with normocapnically raised controls. The ventilatory and blood gas and pH response to CO2 were compared in chronically cannulated test and control rats. Hematocrits and gas pocket CO2 and O2 tensions of mice and rat ventilatory and arterial blood CO2 and O2 tensions and pH responses were not different in test and control groups. There appears to be little or no developmental affect of CO2 suggesting that the reduced CO2 response seen in burrowers is genetically determined.

Neuromuscular organization and regional EMG activity of the pectoralis in the pigeon

In order to improve our understanding of the neuromuscular control of the most massive avian flight muscle, we studied the innervation pattern of the pigeon pectoralis. Nine primary branches from the rostral trunk and nine to ten branches from the caudal trunk of the pectoral nerve were identified by microdissection in ten pigeons. The region of muscle that each branch innervates was delineated by nerve stimulation studies (ten pigeons) and six regions were confirmed by glycogen depletion (ten pigeons). In pigeons, branches from the rostral nerve innervate the anterior 3/5 of the sternobrachialis (SB) head of the pectoralis and branches from the caudal trunk innervate the posterior 1/2 of the SB and all of the throacobrachials (TB). In the SB, individual branches of the rostral pectoral nerve innervate wedge‐shaped muscle regions (each approximately 1.3 cm wide), collectively forming a fan shaped arrangement along the sternal carina. Adjacent muscle regions partially overlap at their boundaries. Within the thoracobrachialis (TB) head of the pectrolis, muscle regions are wider. There is a region in mid‐SB‐where the innervation territories of the rostral and caudal nerves oferlap. Electromyographic (EMG) activity patterns were recorded within ten of the identified muscle regions during take‐off, level flapping flight, and landing. Onset of EMG activity and EMG intensity within various muscle regions exhibits significant differences both within a wingbeat cycle and among different modes of flight. The innervation pattern of the pectoralis presents the anatomical substrate for neuromuscular compartmentalization and differential EMG activity within the pectoralis may reflect sensory‐motor partitioning. The extent to which the neuromuscular compartmentalization of the pectoralis corresponds to its ability to produce an array of force vectors to the wing awaits further more detailed biomechanical studies.

Cardiorespiratory responses of the woodchuck and porcupine to CO2 and hypoxia.

SummaryThe burrow-dwelling woodchuck (Marmota monax) (mean body wt.=4.45±1 kg) was compared to a similar-sized (5.87±1.5 kg) but arboreal rodent, the porcupine (Erithrizon dorsatum), in terms of its ventilatory and heart rate responses to hypoxia and hypercapnia, and its blood characteristics.VT,f,TI andTE were measured by whole-body plethysmography in four awake individuals of each species. The woodchuck has a longerTE/TTOT (0.76±0.03) than the porcupine (0.61±0.03). The woodchuck had a higher threshold and significantly smaller slope to its CO2 ventilatory response compared to the porcupine, but showed no difference in its hypoxic ventilatory response. The woodchuck P50 of 27.8 was hardly different from the porcupine value of 30.7, but the Bohr factor, −0.72, was greater than the porcupines, −0.413. The woodchuck breathing air has PaCO2=48 (±2) torr, PaO2=72 (±6), pHa=7.357 (±0.01); the porcupine blood gases are PaCO2=34.6 (±2.8), PaO2=94.9 (±5), pHa=7.419 (±0.03), suggesting a difference in PaCO2/pH set points. The woodchuck exhibited no reduction in heart rate with hypoxia, nor did it have the low normoxic heart rate observed in other burrowing mammals.

Ventilatory, cardiovascular and metabolic responses to hypoxia and hypercapnia in the armadillo

Armadillos have a low resting metabolic rate and high hemoglobin affinity for their size, a rigid carapace and a semi-fossorial life style. These characteristics could contribute to unusual respiratory responses to hypoxia and hypercapnia which were investigated in this study. Ventilatory and oxygen consumption responses of six adult unanesthetized armadillos to 15, 12, 10 and 8% O2 and 1.5, 3, 5 and 7% CO2 were measured by barometric plethysmography and flow-through respirometry. A significant increase in ventilation occurred in response to 10 and 8% O2 but a decline in oxygen consumption only occurred at 8% inspired O2. The convection requirement response has a threshold at a PaO2 of approximately = 28 Torr which corresponds to a Hb saturation of approximately 70%. Ventilation increased in response to 3% and higher levels of CO2, with no change in oxygen consumption. The magnitude of the ventilatory response to CO2 was similar to other semi-fossorial mammals and less than that of nonburrowing species. However, the pattern of the response was unique in being largely a frequency response with little change in tidal volume, contrary to the tidal volume dominated response to hypercapnia typical of mammals. This feature, not shared by another Xenarthran, the sloth, who lacks a carapace, is likely attributable to the low respiratory system compliance and increased airway resistance resulting from the rigid carapace and small lungs of armadillos and emphasizes the importance of respiratory mechanics in determining breathing pattern.

How stiff is the armadillo? A comparison with the allometrics of mammalian respiratory mechanics

Static respiratory mechanics were examined in the armadillo (Dasypus novemcinctus) and compared with allometric relationships newly derived for adult mammals from values in the literature. Normalised by body weight, chest wall compliance (Cw) in the armadillo is lower than predicted. Lung compliance (Cl) is also low in the armadillo, however it is appropriately matched to the resting lung volume (Vr) (ie. Cl/Vr infinity Mass0.0) and the ratio of Cw/Cl is appropriate for the size of the animal. Respiratory system resistance is high in the armadillo, presumably because of smaller airways associated with the small lung. The power of breathing in the armadillo is comparatively high, mainly due to the high resistive forces. Indeed, the oxidative cost of breathing is approximately double that of a mammal with similar Vr. Hypoxia or hypercapnia are known to invoke an attenuated ventilatory response in the armadillo and one that relies more on changes in frequency rather than volume. While such a breathing pattern helps to reduce the power of breathing it also compromises the degree of hyperventilation achieved.

Scaling of hypercapnic ventilatory responsiveness in birds and mammals.

The possible relationship between CO2 responsiveness and body mass in birds was explored using newly acquired ventilatory data from the barn swallow, Hirundo rustica, and the pigeon, Columbia livia, and that from the literature on four other species. Ventilatory responsiveness (% delta V) of birds to 5% inspired CO2 is scaled to body mass to the 0.145 power (% delta V alpha Mb 0.145). A similar allometric relationship exists for data on 7 species of eutherian mammals taken from the literature (% delta V alpha Mb0.130). The The reduced responsiveness to CO2 in small birds and mammals may be related to an elevated hypoxic ventilatory sensitivity, as demonstrated in mammals (Boggs and Tenney, Respir. Physiol. 58: 245-251, 1984). These scaling relationships may reflect a mechanism for minimizing the inhibition of ventilation resulting from excessive loss of CO2 which thereby permits a higher hypoxic ventilatory response in small species. Other mechanisms, however, could include size related differences in mechanics or alveolar ventilation.

Blood characteristics, tracheal volume and heart mass of burrowing owls (Athene cunicularia) and bobwhite (Colinus virginianus)

1. Measurements of certain hematological and morphological characteristics were made in burrowing owls and bobwhite in search of features that could be associated with the previously described ventilatory adaptations of the burrowing owl to hypoxia and hypercarbia. 2. Values for burrowing owls and bobwhite, respectively, were: P50 = 44.9, 46.0 torr; Hct = 36.6, 38.8 vol%, Hb = 12, 12.3 (g/100 ml); RBC = 2.72 X 10(6), 3.02 X 10(6); log P50/pH = -0.412, -0.485; delta log PCO2/delta pH = -1.39, -1.585. 3. The owls had greater heart weights and smaller tracheal volumes than the bobwhite or the predicted value. 4. No hematological characteristics of the burrowing bird distinguish it from the non-burrowing bird.