RESUMO
The anatomy of the avian lower respiratory system includes a complex interaction between air-filled pulmonary tissues, pulmonary air sacs, and much of the postcranial skeleton. Hypotheses related to the function and phylogenetic provenance of these respiratory structures have been posed based on extensive interspecific descriptions for an array of taxa. By contrast, intraspecific descriptions of anatomical variation for these features are much more limited, particularly for skeletal pneumatization, and are essential to establish a baseline for evaluating interspecific variation. To address this issue, we collected micro-computed tomography (µCT) scans of live and deceased African grey parrots (Psittacus erithacus) to assess variation in the arrangement of the lungs, the air sacs, and their respective invasion of the postcranial skeleton via pneumatic foramina. Analysis reveals that the two pairs of caudalmost air sacs vary in size and arrangement, often exhibiting an asymmetric morphology. Further, locations of the pneumatic foramina are more variable for midline, non-costal skeletal elements when compared to other pneumatized bones. These findings indicate a need to better understand contributing factors to variation in avian postcranial respiratory anatomy that can inform future intraspecific and interspecific comparisons.
RESUMO
The subpectoral diverticulum (SPD) is an extension of the respiratory system in birds that is located between the primary muscles responsible for flapping the wing1,2. Here we survey the pulmonary apparatus in 68 avian species, and show that the SPD was present in virtually all of the soaring taxa investigated but absent in non-soarers. We find that this structure evolved independently with soaring flight at least seven times, which indicates that the diverticulum might have a functional and adaptive relationship with this flight style. Using the soaring hawks Buteo jamaicensis and Buteo swainsoni as models, we show that the SPD is not integral for ventilation, that an inflated SPD can increase the moment arm of cranial parts of the pectoralis, and that pectoralis muscle fascicles are significantly shorter in soaring hawks than in non-soaring birds. This coupling of an SPD-mediated increase in pectoralis leverage with force-specialized muscle architecture produces a pneumatic system that is adapted for the isometric contractile conditions expected in soaring flight. The discovery of a mechanical role for the respiratory system in avian locomotion underscores the functional complexity and heterogeneity of this organ system, and suggests that pulmonary diverticula are likely to have other undiscovered secondary functions. These data provide a mechanistic explanation for the repeated appearance of the SPD in soaring lineages and show that the respiratory system can be co-opted to provide biomechanical solutions to the challenges of flight and thereby influence the evolution of avian volancy.
Assuntos
Voo Animal , Falcões , Respiração , Sistema Respiratório , Asas de Animais , Animais , Evolução Biológica , Fenômenos Biomecânicos/fisiologia , Voo Animal/fisiologia , Falcões/anatomia & histologia , Falcões/classificação , Falcões/fisiologia , Pulmão/anatomia & histologia , Pulmão/fisiologia , Modelos Biológicos , Músculo Esquelético/anatomia & histologia , Músculo Esquelético/fisiologia , Sistema Respiratório/anatomia & histologia , Asas de Animais/fisiologia , Asas de Animais/anatomia & histologia , Masculino , FemininoRESUMO
The vertebrate respiratory system is challenging to study. The complex relationship between the lungs and adjacent tissues, the vast structural diversity of the respiratory system both within individuals and between taxa, its mobility (or immobility) and distensibility, and the difficulty of quantifying and visualizing functionally important internal negative spaces have all impeded descriptive, functional, and comparative research. As a result, there is a relative paucity of three-dimensional anatomical information on this organ system in all vertebrate groups (including humans) relative to other regions of the body. We present some of the challenges associated with evaluating and visualizing the vertebrate respiratory system using computed and micro-computed tomography and its subsequent digital segmentation. We discuss common mistakes to avoid when imaging deceased and live specimens and various methods for merging manual and threshold-based segmentation approaches to visualize pulmonary tissues across a broad range of vertebrate taxa, with a particular focus on sauropsids (reptiles and birds). We also address some of the recent work in comparative evolutionary morphology and medicine that have used these techniques to visualize respiratory tissues. Finally, we provide a clinical study on COVID-19 in humans in which we apply modeling methods to visualize and quantify pulmonary infection in the lungs of human patients.