Background: No integrated comprehensive description of the ultrastructure of the parabronchial epithelium is available. The origin, discharge, and occurrence of the trilaminar substance have not yet been sufficiently studied. Therefore, the main objectives were to classify the cell types of the parabronchial epithelium and to describe their role in manufacturing the trilaminar substance. Methods: Lung tissue of mature quail, domestic fowl, town pigeon, homing pigeon, and barn owl was subjected to standard processing for transmission electron microscopy, both after intratracheal inflation and intravascular perfusion. Results: The atrial epithelium is constituted by granular and squamous atrial cells. Granular cells (1) are confined to the atrial wall; they produce and discharge osmiophilic lamellar bodies. Squamous atrial cells (2) manufacture and discharge a trilaminar substance in sheets sandwiched between the long microvilli emerging from the apical cytoplasm. Their attenuating cell outgrowths overlap granular cells. At the bases of atria, they pass as intermediate squamous atrial cells to the infundibula, contacting the extensions of squamous respiratory cells. The squamous atrial cells undergo distinct structural variations depending on age and environment. Squamous respiratory cells (3) (cellulae squamosae) continuously line the air capillaries and neighboring infundibula. They constitute the epithelial compartment of the blood-gas barrier. The cell bodies extend long, very thin cell outgrowths. The apical surface is smooth and the basal part is invested with a very thin basement membrane. The trilaminar substance originates from granular and agranular endoplas-mic reticulum in the form of convoluted profiles which are discharged as an acellular lining layer on the air surface of squamous respiratory cells. Conclusions: Granular cells are analogous to the type II cells of mamma-lian pulmonary alveolus. Squamous atrial and respiratory cells, of a common embryonic origin, do not meet any counterpart in epithelial cell populations of lung terminal airways in vertebrates. The specific trilami-nar substance-lipoproteinaceous in nature-is a constant compound of atria and air capillaries. Anat.

Avian air capillaries are delicate structures compared to the mammalian pulmonary alveolus. A transmission and scanning electron microscopic study was carried out on several species of birds with the aim of determining the support structures of the avian gas-exchange mantle. Lung tissue of two bird species belonging to strong flying birds (pigeon and barn owl) and two relatively flightless species (domestic fowl and quail) was subjected to standard processing for transmission and scanning electron microscopy after intratracheal inflation. Twisted profiles of lipoproteinaceous trilaminar substance as specific secretory product of avian squamous respiratory cells can be seen in the cell body and cytoplasmic extensions that are wedged between the blood capillaries, partly surrounding them. The intracytoplasmatically located trilaminar complexes form a three-dimensional intricate spiderweb-like system between the blood capillaries and air capillaries, which presumably function as an anchoring and support structure of the gas-exchange tissue. This system is strengthened by retinacula – pairs of attenuated parallel processes of squamous respiratory cells that project to the airway lumen – expanding and bridging the opposite side of air capillaries. The trilaminar substance is discharged in the form of a 15-nm-thick acellular lining layer which is uniquely adapted to the extremely thin respiratory epithelium. The trilaminar substance arises in the cytoplasm of squamous respiratory cells from profiles of granular and smooth endoplasmic reticulum. The integrity and stability of the gas-exchange tissue is likely to be guaranteed by a specific arrangement of the squamous respiratory cells, in which the trilaminar substance plays a paramount role. This general pattern can be observed in strong flying bird species as in the relatively flightless birds.

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The present results demonstrate that avian respiratory macrophages are predominantly located in atrial connective tissue compartments and do not seem to migrate to the airway surfaces.

Summary:A combined scanning electron (SEM) and transmission electron microscopy (TEM) investigation was undertaken to gain insight into the complex structural pattern of the atrial compartment and the gas exchange tissue of parabronchial units in quail and town pigeons. The aim was also to depict the changes taking place in the parabronchial unit in the late prehatching and early posthatching periods in quail. The standard SEM and TEM investigation was carried out in 13 mature quail and 8 town pigeons. The developmental study involved embryonic quail (Days 15, 16, 17), newly hatched quail, quail 24 h after hatching, and quail aged 2, 10, 19, and 25 days (3 individuals per group). The luminal relief of the parabronchus is formed by anastomosing interatrial septa delineating the atrial pits, which are thinner and shallower in pigeons. The atrial bottom opens in mature individuals into 3-6 infundibula. The extracellular material represented by trilaminar substance, which does not appear until hatching, veils the surface relief of the parabronchial epithelium, which is consequently hardly accessible to three-dimensional visualization. Only in town pigeons with fewer discontinuous layers of extracellular material was it possible to visualize the surface of the atrial epithelium, that is, of the granular and squamous atrial cells. The SEM analysis has convincingly shown the intricate spatial organization of atria, infundibula, and air and blood capillaries of the gas exchange tissue. The retinacula, that is, parallelly arranged processes of squamous respiratory cells bridging the air-capillary lumina, were evidenced by SEM and TEM. The complex structure of the avian parabronchus has been successfully demonstrated in the present SEM and TEM study.