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Our department benefits from being part of a vibrant world-class university that holds expansive libraries, archives, and museums, enabling pioneering work in the humanities, arts and sciences. Why German? Beyond language skills, our majors receive a strong liberal arts education that opens doors to law school, graduate study, and international careers in science, business, and IT, at home and in German-speaking countries. Graduates from our prestigious PhD program teach at universities and colleges across the country and abroad.


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This multidisciplinary course explores the history, theory, and international influence of the Bauhaus movement which started as a radical art and design school in Germany in , one hundred years ago. This course explores German popular culture from the last 30 years through various media, including literature, music, performance art, blogs, and movies. Sangamentos on Congo Square? Skip to main content. Department of German Founded in , our department is known for innovative cross-disciplinary research and a dynamic intellectual atmosphere.

Featured Courses. German 3 Special Topics. More details. Staff This course explores German popular culture from the last 30 years through various media, including literature, music, performance art, blogs, and movies. More Featured Courses. Departmental News. New book chapter by Prof.

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Jeroen Dewulf Sangamentos on Congo Square? Increased movement forces a greater degree of compression and expansion of its body cavities, and in turn inflates and deflates more of its air sacs. This not only forces more air through the lungs, but also makes the bird's relative weight lighter. When a bird takes off for flight, the exaggerated movement of its wings creates an air current which fills its air sacs, including those within its bones, and makes the bird light enough to fly.

The abdominal muscles are largely responsible for breathing while at rest. A bird can also use its air sacs to sing by forcing air through its vocal organs like a bagpipe. Some birds can sing while they fly! This is due in part to the bird's ability to sing during inspiration as well as expiration like whistling , as well as an incredible degree of muscle control.

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The exchange of gases simple diffusion of O 2 and CO 2 occurs only between blood capillaries and air capillaries. As air moves through a parabronchus and each successive air capillary, the partial pressure of oxygen P O2 declines as indicated by the decreased density of the stippling because oxygen is diffusing into the blood capillaries associated with each air capillary.


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As a result of this diffusion, the partial pressure of oxygen in the blood leaving the lungs pulmonary vein is higher than that in blood entering the lungs pulmonary artery as indicated by the increased density of the stippling. Relative partial pressures of O 2 and CO 2 1 for air entering a parabronchus initial-parabronchial, P I and air leaving a parabronchus end-parabronchial, P E , and 2 for blood before entering blood capillaries in the lungs pulmonary artery, P A and for blood after leaving the blood capillaries in the lungs pulmonary vein, P V. The partial pressure of oxygen P O2 of venous blood P V is derived from a mixture of all serial air capillary-blood capillary units.

Because of this cross-current exchange the partial pressure of oxygen in avian pulmonary veins P V is greater than that of the air leaving the parabronchus P E ; air that will be exhaled. In mammals, the partial pressure of oxygen in veins leaving the lungs cannot exceed that of exhaled air end-expiratory gas, or P E Figure adapted from Scheid and Piiper Importantly, the partial pressure of oxygen in blood leaving the avian lung is the result of 'mixing'; blood from a series of capillaries associated with successive air capillaries along the length of a parabronchus is mixed as the blood leaves the capillaries and enters small veins.

As a result, the direction of air flow through a parabronchus does not effect the efficiency of the cross-current exchange because gases are only exchanged between blood capillaries and air capillaries, not between the parabronchus and the blood.

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So, in above diagram, reversing the direction of air flow would obviously mean that the air capillary on the far left would have the highest partial pressure of oxygen rather than the air capillary on the far right so the stippling pattern that indicates the amount of oxygen in each air capillary would be reversed. However, because of the 'mixing' of blood just mentioned, this reversal would have little effect on the P V , the partial pressure of oxygen in blood leaving via pulmonary veins the P O2 would likely be a bit lower because some oxygen would have been lost the first time air passed through the neopulmonic parabronchi.

This is important because most birds have neopulmonic parabronchi as well as paleopulmonic parabronchi and, although air flow through paleopulmonic parabronchi is unidirectional, air flow through neopulmonic parabronchi is bidirectional. Diagram showing the flow of air from the parabronchial lumen PL into the air capillaries not shown and arterial blood from the periphery of the parabronchus into the area of gas exchange exchange tissue, ET.

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The orientation between the flow of air along the parabronchus and that of blood into the exchange tissue ET from the periphery is perpendicular or cross-current dashed arrows. The exchange tissue is supplied with arterial blood by interparabronchial arteries IPA that give rise to arterioles stars that terminate in blood capillaries. After passing through the capillaries, blood flows into the intraparabronchial venules asterisks that drain into interparabronchial veins IPV. These in turn empty into the pulmonary vein which returns the blood to the heart.

Maina and Woodward Ventilation and respiratory rate are regulated to meet the demands imposed by changes in metabolic activity e. There is likely a central respiratory control center in the avian brain, but this has not been unequivocally demonstrated. As in mammals, the central control area appears to be located in the pons and medulla oblongata with facilitation and inhibition coming from higher regions of the brain.

It also appears that the chemical drive on respiratory frequency and inspiratory and expiratory duration depend on feedback from receptors in the lung as well as on extrapulmonary chemoreceptors, mechanoreceptors, and thermoreceptors Ludders Central chemoreceptors affect ventilation in response to changes in arterial P CO 2 and hydrogen ion concentration. Peripheral extrapulmonary chemoreceptors, specifically the carotid bodies located in the carotid arteries , are influenced by P O 2 and increase their discharge rate as P O 2 decreases, thus increasing ventilation; they decrease their rate of discharge as P O 2 increases or P CO 2 decreases.

These responses are the same as those observed in mammals. Unlike mammals, birds have a unique group of peripheral receptors located in the lung called intrapulmonary chemoreceptors IPC that are acutely sensitive to carbon dioxide and insensitive to hypoxia. The IPC affect rate and volume of breathing on a breath-to-breath basis by acting as the afferent limb of an inspiratory-inhibitory reflex that is sensitive to the timing, rate, and extent of CO 2 washout from the lung during inspiration Ludders Pulmonary surfactant in birds: From birds to humans: The avian respiratory system: Environ Health Perspectives Activity of three muscles associated with the uncinate processes of the giant Canada Goose Branta canadensis maximus.

Journal of Experimental Biology Evolution of surface activity related functions of vertebrate pulmonary surfactant. Clin Exp Pharmacol Physiol. Quantitative analysis of the respiratory system of the House Sparrow, Budgerigar, and Violet-eared Hummingbird. The lung air sac system of birds. Advances in Anatomy, Embryology, and Cell Biology A comparative perspective of the progressive integration of respiratory system, locomotor apparatus and ontogenetic development.

Function of intracoelomic septa in lung ventilation of amniotes: Physiological and Biochemical Zoology Inhaled anesthesia for birds. Recent advances in veterinary anesthesia and analgesia: The morphometry of the avian lung. Themes and principles in the design and construction of the gas exchangers. Structure, function and evolution of the gas exchangers: Journal of Anatomy Functional morphology of the avian respiratory system, the lung-air system: The lung of the Emu, Dromaius novaehollandiae: An allometric study of the pulmonary morphometric parameters in birds, with mammalian comparison.

The bioengineering dilemma in the structural and functional design of the blood-gas barrier.


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Three-dimensional serial section computer reconstruction of the arrangement of the structural components of the parabronchus of the Ostrich, Struthio camelus lung.