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Extension of oxygen tolerance in man: philosophy and significance.

The respiration of oxygen over a range of partial pressures higher than in the natural environment has expanding usefulness in health and disease. It provides for denitrogenation of the astronaut to prevent aerospace decompression sickness, it facilitates diving of many forms and improves safety in decompression after diving, and it is the key to therapy of diving and iatrogenic gas embolic diseases. In the continuum of general and hyperbaric medicine, it is essential for sustaining viability of damaged or diseased tissues not adequately oxygenated at natural oxygen pressures. Over the entire range of its clinical and operational usefulness, the pressure and duration of tolerable exposure to oxygen is limited by adverse effects on multiple chemical targets, cells, tissues, and organ functions. The rates of development and the qualitative expressions of these effects are different at different respired oxygen pressures. Successful extension of oxygen tolerance, as by slowing the rate of development of adverse effects, will further expand the medical and operational usefulness and safety of oxygen in normal, hypobaric, and hyperbaric environments. A prerequisite baseline for overall extension of oxygen tolerance is the quantitative investigation of early stages of toxic oxygen effects upon specific chemical and composite functions of multiple organ systems, including rates of development and rates of recovery. The practicality of limited oxygen tolerance extension by systematic interruption of oxygen exposure has been demonstrated and the procedure widely used. Broad present goals are both to establish oxygen tolerance for specific tissues and to optimize tolerance extension over the full range of useful oxygen pressures.

Adaptation, Physiological↗

The effect of Losartan, an angiotensin II antagonist, on cardiac function, mass and morphology in rats after repeated hyperbaric exposures.

Previously, we have found development of left ventricular hypertrophy (LVH), myocardial necrosis and haemodynamic changes in rats after repeated hyperbaric exposures to 5 bar. Exactly the same rat model and experimental set-up was applied in the present study. The purpose of this study was to investigate if angiotensin II (Ang-II) plays a role in the development of these cardiac changes. Losartan, a recently developed non-peptide Ang-II receptor antagonist, was administrated (20 mg kg-1 day-1) to each rat for 40 days. Nine test rats underwent chamber dives daily for 40 consecutive days, and 10 control rats were exposed in the same chamber for an equal period of time, but in air at 1 bar. After 40 days, test rats and control rats had equal right and left ventricular myocardial mass/100 g-1 body mass, thus indicating that Losartan inhibits hyperbarically induced LVH. Microscopic examination revealed no changes in the left ventricle, indicating that Losartan prevents myocardial necrosis. The left ventricular pressure (LVP) and the maximal velocity of LVP increase and decrease (+/-dP/dt) were similar in the test rats compared to the control rats at 1 bar. Previously we found a higher LVP and dP/dt in non-treated test rats in otherwise identical experiments. This indicates that Losartan "normalizes" the cardiac function of test rats after repeated hyperbaric exposures. The systolic arterial pressure, heart rate (HR) and respiratory frequency (RF) were similar in the two groups at 1 bar. However, treatment with Losartan lowered the blood pressure compared to previously non-treated rats. In conclusion, long-term Ang-II receptor blockade prevented previously shown changes in cardiac function and morphology, as well as myocardial mass, after 40 consecutive exposures to 5 bar.

Adaptation, Physiological↗

Physiologic and perceptual responses to hypercarbia during warm- and cold-water immersion.

Thermoregulatory, respiratory, and perceptual responses to acute CO2 exposure during light exercise (75 W) were assessed in 12 U.S. Navy divers clad only in swim trunks while immersed to the neck in water at 18 degrees and 34 degrees C. The CO2 exposures consisted of a linear 10-min ramp increase in the inspired fraction of CO2 (FICO2) from 0 to 6% followed by 5 min of breathing 6% CO2. The ability to detect and rate the severity of hypercarbia, as well as subjective changes in thermal comfort, were assessed by comparing subjective ratings given during the CO2 exposures with those given during immersion trials where the FICO2 was maintained at 0%. Hypercarbia was recognized earlier and, at a given PETCO2, was perceived to be greater during cold- than during warm-water immersions (P < 0.01). The CO2 exposures did not affect the thermal balance of divers as assessed by changes in heat flux and rectal temperature. However, increased feelings of warmth were reported during both the cold and warm immersions when breathing raised concentrations of CO2 (P < 0.01). During the cold immersions, acute exposure to 6% CO2 significantly decreased forearm blood flow (P < 0.05), but did not affect finger blood flow. It is concluded that PETCO2 levels can reach unacceptably high concentrations (> 60 mmHg) before hypercarbia can be reliably detected by working divers. Furthermore, the ability to detect hypercarbia is poorer when immersed in warm water than when in cold water.

Adult↗

[Continuous electrocardiographic recording during a first parachute jump].

The behavior of the cardiac rhythm under intense stress was studied with continuous electrocardiographic recording during the first jump with an automatic parachute in 13 members of the Universidad Nacional Autónoma de México sky diving club. There were 12 male (92.3%) and one female (7.68) with an average age of 22.8 years. A complete clinical history, 12 lead electrocardiogram and a treadmill stress test were performed two weeks before jumping. A two-channel Holter recording system was hooked up 30 min. before boarding the airplane and was turned off one hour later. The heart rate values were compared two weeks prior (64.5 beats/min), before (112.8 beats/min), during (170 beats/min) and after the jump (122.8 beats/min). The mean difference between each phase was statistically significant with p less than 0.001 values. The observed cardiac rhythm was sinus tachycardia in each case. In six cases (43.6%) 22 episodes of sudden decrease of the heart rate were seen and there were no major rhythm or conduction disturbances. The urinary catecholamines were similar in nine cases (69.2%) after the jump (x 51.2 micrograms/100 ml) and two weeks later in four control cases (x 10.3 micrograms/100 ml). We concluded there were no significant rhythm disturbances in the presence of an intense but brief stress condition in young healthy people. This study was classified as longitudinally, descriptive, experimental and projective.

Adult↗

Effects of SCH 42495, a neutral endopeptidase inhibitor, on cardiac function and mass in rats after repeated hyperbaric exposures.

The aim of the present study was to investigate the effects of inhibition of neutral endopeptidase on left ventricular hypertrophy (LVH), myocardial necrosis and haemodynamic changes, previously shown in rats after repeated hyperbaric exposures to 5 bar, by using the novel neutral endopeptidase inhibitor SCH 42495. Ten test rats underwent chamber dives daily for 40 consecutive days, and 10 control rats were exposed in the same chamber for an equivalent period of time, but in air at 1 bar. The rats received SCH 42495 orally (30 mg/kg twice a day) for 40 days. After 40 days, the body mass was identical in the two groups. Test rats and control rats had equal heart mass (both totally and left ventricular myocardium, including the ventricular septum/100 g), thus indicating that long-term treatment with SCH 42495 inhibits hyperbarically induced LVH. The left ventricular pressure (LVP) and the maximal velocity of LVP increase and decrease (+/- dP/dt) were similar in the test rats compared to the control rats at 1 bar. Previously, we found a higher LVP and dP/dt in non-treated test rats in otherwise identical experiments. This indicates that SCH 42495 "normalizes" the cardiac function of test rats after repeated hyperbaric exposures. The systolic arterial pressure, heart rate (HR) and respiratory frequency (RF) were similar in the two groups throughout the experiments. However, treatment with SCH 42495 lowered the blood pressure compared to previously non-treated rats. In conclusion, long-term administration of the neutral endopeptidase inhibitor SCH 42495 prevented previously shown changes in cardiac function as well as myocardial mass after 40 consecutive exposures to 5 bar.

Adaptation, Physiological↗

Rat liver injury after normothermic ischemia is prevented by a phosphinic matrix metalloproteinase inhibitor.

Hepatic ischemia occurs in liver transplantation, hemodynamic or cardiogenic shock, and liver resection associated with trauma or tumor. Ischemia/reperfusion (I/R) injury results in microcirculation failure followed by apoptosis and necrosis. Matrix metalloproteinases (MMPs) are involved in many physiological and pathological processes, but their expression and function during liver I/R remains poorly documented. In this study, we evaluated the expression of nine MMPs and their natural inhibitors, tissue inhibitors of MMPs (TIMPs), in a rat model of liver I/R. Analysis of MMP and TIMP expression show that although most of these genes are not constitutively expressed in the normal liver, they are induced in a specific time-dependent manner following I/R. Stromelysin-1, gelatinase B, and collagenase-3 are induced during the early phase of acute liver injury associated with inflammation and increased necrosis/apoptosis, whereas gelatinase A, membrane type-MMP, stromelysin-3, metalloelastase, TIMP-1, and TIMP-2 are essentially detectable during the recovery phase of liver injury corresponding to hepatocyte regeneration. This observation suggested that MMPs and TIMPs could play both deleterious and beneficial roles following I/R. We thus tested the effect of a specific phosphinic MMP inhibitor on acute liver I/R injury. Inhibition of MMP activity was shown to significantly decrease liver injury in ischemic/reperfused liver tissue as assessed by histological studies and serum hepatic enzyme levels. We therefore propose that MMP inhibitors may be of clinical relevance in liver-associated ischemic diseases or after liver transplantation.

Animals↗

A comparison of cardiovascular and autonomic adjustments to three types of cold stimulation tasks.

Two types of 'cold pressor' tasks are used frequently in research settings: immersing the hand or foot in ice water, and applying an icebag to the forehead. Both tasks have commonly been selected as 'alpha-adrenergic' tasks due to expected increases in blood pressure and peripheral resistance. However, the forehead cooling task has been used by others to produce increased vagal tone due to the elicitation of the 'diving reflex' and subsequent bradycardia. This differs from the prototypical increase in heart rate during hand immersion. The present study directly compared the cardiovascular adjustments of hand immersion and forehead stimulation. As stimulation of the trigeminal nerve is thought to increase vagal activity during forehead cooling, a third condition in which most of the face was covered with an icebag was included to ostensibly stimulate more of the trigeminal. 18 males had counterbalanced exposures to hand immersion, forehead cooling, and facial cooling for 90 s each. Cardiovascular variables derived from impedance cardiography and the ECG were measured. Respiration was paced at 14 breaths/min to facilitate comparisons of respiratory sinus arrhythmia across conditions. No differences among baseline or tasks were found for respiratory rate or cardiac output. The pattern of results for hand immersion was that of increased heart rate with moderate blood pressure increases. In contrast, the forehead and facial cooling tasks elicited small heart rate decreases with little change in blood pressure. Facial cooling elicited significantly more vagal activation than hand cooling as indexed by respiratory sinus arrhythmia. The patterns of response for facial and forehead cooling were almost identical. Probable reasons for the lack of significant blood pressure responses during the facial and forehead cooling are discussed.

Adolescent↗

Inhibition of intracellular esterases by antitumour chloroethylnitrosoureas. Measurement by flow cytometry and correlation with molecular carbamoylation activity.

Antitumour chloroethylnitrosoureas (Cnus) decompose in physiological conditions yielding alkylating species and organic isocyanates. While antitumour activity is mainly attributed to the alkylation of DNA, carbamoylation of intracellular proteins by isocyanates may also have pharmacological and toxicological relevance. We previously reported a novel dynamic flow cytoenzymological assay for esterase inhibition in intact murine cells by BCNU and related isocyanates, and proposed that this might form the basis of an assay for intracellular carbamoylation. We have now examined a wide range of Cnus, isocyanates, and alkylating agents for their ability to inhibit cellular esterases. BCNU, CCNU, their derived isocyanates, and the 4-OH metabolites of CCNU exhibited potent inhibitory activity (I50 values 5.5 x 10(-5)-7.3 x 10(-4) M). Chlorozotocin and GANU were relatively inactive (I50 much greater than 10(-2) M). ACNU, TCNU and the 2-OH metabolites of CCNU exhibited intermediate activity (I50 values 1.1 x 10(-3)-2.3 x 10(-2) M). Compounds not able to form isocyanates were essentially inactive. Poor membrane permeability was also implicated in the weak activity of chlorozotocin and GANU. There was overall a good correlation between esterase inhibition and chemical carbamoylating activity, but some particular differences were identified. We concluded that flow cytoenzymological assay appears to have the potential to provide useful measurement of intracellular protein carbamoylation by existing Cnus and novel derivatives, and also offers the advantage of cell subpopulation identification for in vivo evaluation of these agents.

Animals↗

Physiological responses to head-out immersion in water at 11 ATA.

Cardiorespiratory, thermal, and renal responses to a 30-min head-out immersion in 15 degree C water were studied at 1-ATA air and 11-ATA helium-oxygne environments in four male subjects wearing dry suits. Cardiorespiratory responses to immersion (reductions in heart rate, expiratory reserve volume, vital capacity, and thoracic impedance; and increases in stroke volume, cardiac output, and inspiratory capacity) were comparable at both pressures. However, thermal responses to immersion (a reduction in mean skin temperature and increases in skin heat flux and suit conductance) were significantly greater at 11 ATA compared to those at 1 ATA. The rate of urinary excretion of norepinephrine increased significantly during and after immersion at 11 ATA but not at 1 ATA. In contrast, the urinary excretion of epinephrine was not altered by pressure or immersion. The immersion diuresis was greater and lasted longer at 11 ATA than at 1 ATA although there was no difference in the endogenous creatinine excretion . This diuresis was accompanied by a significant natriuresis which was more marked at 1 ATA than at 11 ATA. At 1 ATA, the urinary excretion of both aldosterone and antidiuretic hormone (ADH) decreased during immersion. At 11 ATA, the rate of excretion of these hormones before immersion was lower compared to that at 1 ATA and did not change significantly during immersion. These results indicate that immersion in a hyperbaric helium-oxygen environment presents a greater cold stress than at 1-ATA air, and also that immersion diuresis and natriuresis at high pressure may be induced by a factor other than inhibition of aldosterone and ADH.

Adult↗

Pulmonary gas exchange is not impaired 24 h after extravehicular activity.

Extravehicular activity (EVA) during spaceflight involves a significant decompression stress. Previous studies have shown an increase in the inhomogeneity of ventilation-perfusion ratio (VA/Q) after some underwater dives, presumably through the embolic effects of venous gas microemboli in the lung. Ground-based chamber studies simulating EVA have shown that venous gas microemboli occur in a large percentage of the subjects undergoing decompression, despite the use of prebreathe protocols to reduce dissolved N(2) in the tissues. We studied eight crewmembers (7 male, 1 female) of the International Space Station who performed 15 EVAs (initial cabin pressure 748 mmHg, final suit pressure either approximately 295 or approximately 220 mmHg depending on the suit used) and who followed the denitrogenation procedures approved for EVA from the International Space Station. The intrabreath VA/Q slope was calculated from the alveolar Po(2) and Pco(2) in a prolonged exhalation maneuver on the day after EVA and compared with measurements made in microgravity on days well separated from the EVA. There were no significant changes in intrabreath VA/Q slope as a result of EVA, although there was a slight increase in metabolic rate and ventilation (approximately 9%) on the day after EVA. Vital capacity and other measures of pulmonary function were largely unaltered by EVA. Because measurements could only be performed on the day after EVA because of logistical constraints, we were unable to determine an acute effect of EVA on VA/Q inequality. The results suggest that current denitrogenation protocols do not result in any major lasting alteration to gas exchange in the lung.

Adaptation, Physiological↗

Ventilatory mechanics and the effects of water depth on breathing pattern in the aquatic caecilian Typhlonectes natans.

The breathing pattern in the aquatic caecilian Typhlonectes natans was investigated by recording airflow via a pneumotachograph under unrestrained normal physiological conditions. Ventilatory mechanics were assessed using airflow and pressure measurements from the buccal cavity and trachea. The breathing pattern consisted of an expiratory phase followed by a series of 10-15 small buccal pumps to inflate the lung, succeeded by a long non-ventilatory period. T. natans separate the expiratory and inspiratory gases in the buccal cavity and take several inspiratory pumps, distinguishing their breathing pattern from that of sarcopterygians. Hydrostatic pressure assisted exhalation. The tracheal pressure was greater than the water pressure at that depth, suggesting that pleuroperitoneal pressure as well as axial or pulmonary smooth muscles may have contributed to the process of exhalation. The frequency of lung ventilation was 6.33+/-0.84 breaths h(-)(1), and ventilation occurred via the nares. Compared with other amphibians, this low ventilatory frequency suggests that T. natans may have acquired very efficient pulmonary respiration as an adaptation for survival in their seasonally fluctuating natural habitat. Their respiratory pathway is quite unique, with the trachea separated into anterior, central and posterior regions. The anterior region serves as an air channel, the central region is attached to the tracheal lung, and the posterior region consists of a bifurcated air channel leading to the left and right posterior lungs. The lungs are narrow, elongated, profusely vascularized and compartmentalized. The posterior lungs extend to approximately two-thirds of the body length. On the basis of their breathing pattern, it appears that caecilians are phylogenetically derived from two-stroke breathers.

Amphibians↗

Patent foramen ovale: a review of associated conditions and the impact of physiological size.

Patent foramen ovale (PFO) is implicated in platypnea-orthodeoxia, stroke and decompression sickness (DCS) in divers and astronauts. However, PFO size in relation to clinical illness is largely unknown since few studies evaluate PFO, either functionally or anatomically. The autopsy incidence of PFO is approximately 27% and 6% for a large defect (0.6 cm to 1.0 cm). A PFO is often associated with atrial septal aneurysm and Chiari network, although these anatomic variations are uncommon. Methodologies for diagnosis and anatomic and functional sizing of a PFO include transthoracic echocardiography (TTE), transesophageal echocardiography (TEE) and transcranial Doppler (TCD), with saline contrast. Saline injection via the right femoral vein appears to have a higher diagnostic yield for PFO than via the right antecubital vein. Saline contrast with TTE using native tissue harmonics or transmitral pulsed wave Doppler have quantitated PFO functional size, while TEE is presently the reference standard. The platypnea-orthodeoxia syndrome is associated with a large resting PFO shunt. Transthoracic echocardiography, TEE and TCD have been used in an attempt to quantitate PFO in patients with cryptogenic stroke. The larger PFOs (approximately > or =4 mm size) or those with significant resting shunts appear to be clinically significant. Approximately two-thirds of divers with unexplained DCS have a PFO that may be responsible and may be related to PFO size. Limited data are available on the incidence of PFO in high altitude aviators with DCS, but there appears to be a relationship. A large decompression stress is associated with extra vehicular activity (EVA) from spacecraft. After four cases of serious DCS in EVA simulations, a resting PFO was detected by contrast TTE in three cases. Patent foramen ovales vary in both anatomical and functional size, and the clinical impact of a particular PFO in various situations (platypnea-orthodeoxia, thromboembolism, DCS in underwater divers, DCS in high-altitude aviators and astronauts) may be different.

Aerospace Medicine↗

Arterial Windkessels in marine mammals.

In marine mammals, the aortic arch is enlarged relative to the descending aorta to varying degrees in different species. The ratio of maximal diameter of the arch to that of the thoracic aorta is about 2.3 in the harbour seal (Phoca vitulina), 3.6 in the Weddell seal (Leptonychotes weddelli) and 3.2 in the fin whale (Balaenoptera physalus), compared with only 1.4 in the dog. This anatomical specialisation probably provides increased volume capacitance in the arterial circulation as an adaptation to diving bradycardia. Data on the morphometric and mechanical properties of aortic tissues from seals and fin whale are compared. In the harbour seal, more than 80% of the volume change in the entire thoracic aorta that results from a pressure pulse occurs in the bulbous arch, and this is more than 90% in the Weddell seal and fin whale. The enhanced capacitance of the arch in the harbour seal is primarily due to its larger diameter, as the relative wall thickness and elasticity of the arch and thoracic aorta are the same. A similar situation appears to exist in the larger Weddell seal, although extrapolation of the pressure-volume curves suggests that the arch might be somewhat less stiff than the thoracic aorta. In addition to being greatly expanded, the aortic arch of the fin whale is also much more distensible than the relatively thin-walled and much stiffer descending aorta. At the estimated mean blood pressure, the elastic modulus of this vessel is 12 MPa, or 30 times that of the aortic arch. The major haemodynamic consequence of this type of arterial modification is that the aortic arch acts as a Windkessel, i.e. the capacitance of the aorta is increased significantly close to the heart, leading to a reduced characteristic impedance and probably reduced pulsatility in the descending aorta. In the extreme case of the whale, the arterial capacitance is shifted entirely to the arch, and the impedance change at the entrance to the thoracic aorta is so high that this probably represents the major reflection site in the arterial tree.

Adaptation, Physiological↗

The opposing physiological effects of high pressures and inert gases.

The physiological effects on mammals of elevated pressures (approximately 100 atmospheres) must be considered in the context of the inert gases breathed. The most striking effect of pressure per se is a central hyperexcitability manifest at first by trembling of the entremities and finally by convulsions. Paralysis and death occur at higher pressures. The primary effects of the inert gases breathed are inert gas narcosis and general anesthesia. The exciting effects of pressure per se and the depressive effects of the inert gases tend to oppose each other. Thus consciousness may be restored to anesthetized mice by raising the pressure, and conversely the threshold pressure that causes convulsions is elevated in the presence of anesthetics. These mutually antagonistic effects can be rationalized in terms of model which proposes that both anesthetics and pressure non-specifically perturb thelipid bilayer regions of neutral membranes. This model is termed the critical volume hypothesis. Anthesthetics dissolve in and expand these lipid bilayer regions, while pressure causes mechanical compression. Expansion leads to anesthesia and compression to convulsions if a critical degree of change is achieved. At elevated partial pressures of inert gas the gas-induced expansion is opposed by the compression of pressure per se. With very insoluble gases, such as helium, this expansion is so small that net compression results and the effects of helium differ little from those of pressure per se. With more soluble gases, such as nitrogen, net expansion results in inert gas narcosis and anesthesia. The critical volume hypothesis enables "safe" mixtures of "expanding" and "compressing" gases to be defined. These enable higher pressures to be better tolerated by mammals.

Anesthesia↗

Cold tolerance in mammalian cells.

As whole organisms, most mammals have a poor tolerance for hypothermia. But their cells may have a capacity for a far wider cold tolerance, which may be expressed in peripheral tissues, sporadically in core tissue and in cultured cells. Against this background the cold resistance of cells of deep hibernators may be seen as the extreme of a continuum and is complicated by the consideration that the voluntary hypothermia of hibernation is probably in most cases a metabolic adaptation to forestall starvation. Similarly, cold resistance of peripheral tissues may in diving animals be confounded by the need to be adapted to hypoxia as well. Hence, attempts to analyse cold resistance by comparisons of absolute rates of arbitrarily chosen reactions may be misleading. A more useful approach is analysis of maintenance of balance: balance between ATP synthesis and utilization, balance between macromolecule synthesis and degradation and balance between pumps and leaks. Cation pumps and leaks constitute a major component of energy utilization and are central to other cell functions, even during minimal metabolism. Hence, the maintenance of ion gradients is a central issue in understanding adaptation not only to hypothermia but also to starvation and hypothermia. Of the three hypometabolic states, hypothermia has been best studied in this regard. In most cases, passive permeability is more reduced at low temperature in cold-tolerant cells than in cold-sensitive ones. In some cases there is also a difference in Na-K pump activity and perhaps in ATP dependent Ca-pump activity. Pump activities and probably the maintenance of minimal leak require ongoing metabolism. The question of whether, in cold-sensitive cells, energy supplies are adequate at low temperature was once the focus of this field, but has been ignored for a decade without having been fully resolved. There are many instances of less temperature sensitivity of specific metabolic activities (mitochondrial respiration, etc.) in hibernators than in non-hibernators, without any verification of whether this is essential for survival at low body temperature. Certainly, robust pumping has been found in some failing cold-sensitive cells at low temperature, suggesting no shortage of ATP in these cases, but in other cases the issue may be a more complex one than just that of ATP availability.(ABSTRACT TRUNCATED AT 400 WORDS)

Adaptation, Physiological↗

[Morphologic-functional study of the locomotor system of penguins as a general model of movement in under-water flight. I].

Regarding several theories of the evolution of the Sphenisciformes the specific morpho-physiological alterations for the changeover from aerial to underwater life are discussed. The peculiarities in the Penguin's "construction" become comprehensible as strong adjustments to the subaquatic locomotion. Surely they took their origin from the equipment of flying birds. The present data of the kinematics of the underwater locomotion show, that propulsion is produced in the same principal way by the flapping wings as in aerial flight. Therefore the short term "underwater flight" for the Penguin's style of locomotion is justified. Known data of swimming performance suggest that its essential adaptation is not that to top achievements but more to an economical use of energy budget. The favourable hydrodynamic characteristics of the Penguin body may be well interpreted from this point of view. The peculiarity of underwater flight is the absence of the necessity to produce a weight-compensating force. In order to create thrust forces in an appropriate magnitude during up- and downstroke of the beating cycle the upstroke must be powered. The anatomical architecture and the mode of operation of the parts of the muscle system must be adjusted to this demand. Based on these statements, the anatomy of active and passive apparatus of movement was studied by dissection of 26 individuals of Pygoscelis papua, P. antarctica, P. adeliae, Eudyptes chrysolophus, and Aptenodytes forsteri. Besides the functional explanation of the Articulatio sternocoracoidea (diverging considerably from the usual type in birds), a new interpretation is given for the structures of the Articulatio humeri. In this context, the role of the Ligamentum acrocoracohumerale as an important element for coordination of the motion processes in the shoulder joint is elucidated. The essential curvature of the Caput humeri is found to be satisfactorily approximated by a logarithmic spiral. The understanding of the mechanics of bones and tendons leads to a reinterpretation of the role of several groups of muscles which is described in detail. Besides of the preponderant thrust producing (flapping) muscles working mainly in the isotonic manner, muscles can be distinguished which are managing the transfer of the produced forces to the body operating thereby in the isometric way. Another group of muscles has to control the position of the humerus adjusting in this way the hydrodynamic angle of attack corresponding to the respective flow conditions.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗