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Ventilation and the air sacs
Unlike mammals, birds do not possess a muscular diaphragm, and the lungs do not act as a pair of bellows. The design of the avian respiratory system is extraordinarily efficient, so that birds can extract sufficient oxygen for life even while undertaking strenuous exercise at altitudes of up to 7 or even 8 Km.
While the relative weight of the lungs is comparable to that seen in mammals, their relative volume is only one-tenth. This is because air flows through the lungs in a constant one-way stream, unlike mammals and reptiles where there is an inefficient tidal ebb and flow.
In addition the exchange surface (ie distance from air to blood) is much thinner in birds.
A major feature of the respiratory system of birds is pneumatisation of the long bones, the vertebral column and even the skull. These air spaces connect with the air sacs (below).
The air flow is a complex circulation involving pumping action of thin-walled air sacs in the thorax and abdomen. These extend into the long bones: in the 18th century John Hunter showed that birds could still breath with a blocked windpipe provided one of the long bones (he tried both the femur and the humerus) was connected to the outside air. Most of the major bones of the body connect with the air sac system. This gives lightness as well as being a reservoir of oxygen.
The major features of the respiratory system are as follows. (See Fig below )
The nostrils are generally at the base of the upper mandible. There is no soft palate and both the oropharynx and the choanal opening pass air to the glottis to enter the larynx.
The trachea connects this to the syrinx, the site of vocalization generally at the tracheal bifurcation. This has three potential vibrating surfaces, which explains why birds like Magpies can sing complex harmonies of different pitch simultaneously. Birds vary considerably in the musculature and anatomy of the syrinx and this of course reflects the potential complexity of their vocalisation.
The primary bronchi connect the syrinx to each lung, but the air passes through in the mesobronchi to the abdominal and posterior (caudal) thoracic air sacs. These lie ventro-lateral to the abdominal viscera: between the intestines and the abdominal wall. In addition there are paired anterior air sacs and a single interclavicular air sacarising from the mesobronchi, but there is considerable variation on this basic pattern between species.
From the caudal air sacs the air now flows cranially through the lungs by dorsobronchi and ventrobronchi. These branch out into a complex arrangement of air capillaries or parabronchi. Blood capillaries in the walls of these flow counter to the air stream, allowing for a highly efficient gas exchange.
The stale air then passes to the anterior air sacs and to an unpaired interclavicular air sac, whence it discharges via the trachea.
There is thus a four-beat cycle to respiration:--
(1) First inhalation. Air passes to the posterior air sacs, with expansion of the abdomen.
(2) First exhalation. The abdomen contracts forcing air through the lungs.
(3) Second inhalation. As the abdomen expands again the stale air in the lungs is forced out to the anterior sacs.
(4) Second exhalation. Contraction of the abdomen and anterior sacs drives stale air out of the trachea while fresh air floods into the exchange region.
Most birds have 9 air sacs:
one interclavicular sac
two cervical sacs
two anterior thoracic sacs
two posterior thoracic sacs
two abdominal sacs
Functionally, these 9 air sacs can be divided into anterior sacs (interclavicular, cervicals, & anterior thoracics) & posterior sacs (posterior thoracics & abdominals). Air sacs have very thin walls with few blood vessels. So, they do not play a direct role in gas exchange. Rather, they act as a 'bellows' to ventilate the lungs (Powell 2000).
Source:
http://numbat.murdoch.edu.au/Anatomy/avian/fig3.2.GIF
Air sacs and axial pneumatization in an extant avian. The body of bird in left lateral view, showing the cervical (C), interclavicular (I), anterior thoracic (AT), posterior thoracic (PT), and abdominal (AB) air sacs. The hatched area shows the volume change during exhalation. The cervical and anterior thoracic vertebrae are pneumatized by diverticula of the cervical air sacs. The posterior thoracic vertebrae and synsacrum are pneumatized by the abdominal air sacs in most taxa. Diverticula of the abdominal air sacs usually invade the vertebral column at several points. Diverticula often unite when they come into contact, producing a system of continuous vertebral airways extending from the third cervical vertebra to the end of the synsacrum. Modified from Duncker 1971 (Wedel 2003).
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