Biomedical subjects

J N Maina

Publications and source records attributed to J N Maina.

At least 19 recordsLinked to original sources

PubMed2006

Spectacularly robust! Tensegrity principle explains the mechanical strength of the avian lung.

Among the air-breathing vertebrates, the respiratory system of birds, the lung-air sac system, is remarkably complex and singularly efficient. The most perplexing structural property of the avian lung pertains to its exceptional mechanical strength, especially that of the minuscule terminal respiratory units, the air- and the blood capillaries. In different species of birds, the air capillaries range in diameter from 3 to 20 micro m: the blood capillaries are in all cases relatively smaller. Over and above their capacity to withstand enormous surface tension forces at the air-tissue interface, the air capillaries resist mechanical compression (parabronchial distending pressure) as high as 20 cm H(2)O (2 kPa). The blood capillaries tolerate a pulmonary arterial vascular pressure of 24.1 mmHg (3.2 kPa) and vascular resistance of 22.5 mmHg (3 kPa) without distending. The design of the avian respiratory system fundamentally stems from the rigidity (strength) of the lung. The gas exchanger (the lung) is uncoupled from the ventilator (the air sacs), allowing the lung (the paleopulmonic parabronchi) to be ventilated continuously and unidirectionally by synchronized bellows like action of the air sacs. Since during the ventilation of the lung the air capillaries do not have to be distended (dilated), i.e., surface tension force does not have to be overcome (as would be the case if the lung was compliant), extremely intense subdivision of the exchange tissue was possible. Minuscule terminal respiratory units developed, producing a vast respiratory surface area in a limited lung volume. I make a case that a firm (rigid) rib cage, a lung tightly held by the ribs and the horizontal septum, a lung directly attached to the trunk, specially formed and compactly arranged parabronchi, intertwined atrial muscles, and tightly set air capillaries and blood capillaries form an integrated hierarchy of discrete network system of tension and compression, a tensegrity (tensional integrity) array, which absorbs, transmits, and dissipates stress, stabilizing (strengthening) the lung and its various structural components.

Animals↗

PubMed2006

The morphology of the pecten oculi of the ostrich, Struthio camelus.

The pecten oculi is a structure peculiar to the avian eye. Three morphological types of pecten oculi are recognized: conical type, vaned type and pleated type. The pleated type has been well studied. However, there exists only scanty data on the morphology of the latter two types of pectens. The structure of the vaned type of pecten of the ostrich, Struthio camelus was investigated with light and electron microscope. The pecten of this species consists of a vertical primary lamella that arises from the optic disc and supports 16-19 laterally located secondary lamellae, which run from the base and confluence at the apex. Some of the secondary lamellae give rise to 2 or 3 tertiary lamellae. The lamellae provide a wide surface, which supports 2-3 Layers of blood capillaries. Pigmentation is highest at the distal ends of the secondary and tertiary Lamella where blood capillaries are concentrated and very scanty on the primary and the proximal ends of the secondary lamella where the presence of capillaries is much reduced. In contrast to the capillaries of the pleated pecten, the endothelium of the capillaries in the pecten of the ostrich exhibits very few microvilli. These observations suggest that the morphology of the pecten of the ostrich, a flightless ratite bird is unique to the pleated pecten and is designed to meet the balance between optimal vision and large surface area for blood supply and yet ensuring it is kept firmly erect within the vitreous.

Animals↗

PubMed2004

Morphogenesis of the laminated, tripartite cytoarchitectural design of the blood-gas barrier of the avian lung: a systematic electron microscopic study on the domestic fowl, Gallus gallus variant domesticus.

Formation of a thin blood-gas barrier in the respiratory (gas exchange) tissue of the lung of the domestic fowl, Gallus gallus variant domesticus commences on day 18 of embryogenesis. Developing from infundibulae, air capillaries radiate outwards into the surrounding mesenchymal (periparabronchial) tissue, progressively separating and interdigitating with the blood capillaries. Thinning of the blood-gas barrier occurs by growth and extension of the air capillaries and by extensive disintegration of mesenchymal cells that constitute transient septa that divide the lengthening and anastomosing air capillaries. After they contact, the epithelial and endothelial cells deposit intercellular matrix that cements them back-to-back. At hatching (day 21), with a thin blood-gas barrier and a large respiratory surface area, the lung is well prepared for gas exchange. In sites where air capillaries lie adjacent to each other, epithelial cells contact directly: intercellular matrix is lacking.

Animals↗

PubMed2004

Systematic analysis of hematopoietic, vasculogenetic, and angiogenetic phases in the developing embryonic avian lung, Gallus gallus variant domesticus.

In the embryonic lung of the domestic fowl, Gallus gallus variant domesticus, hematogenetic and vasculogenetic cells become ultrastructurally clear from day 4 of development. In the former group of cells, filopodial extensions coalesce, cytoplasm thickens, and accumulating hemoglobin displaces the nucleus peripherally while in the latter, conspicuous filopodial extensions and large nuclei develop as the cells assume a rather stellate appearance. From day 5, erythrocytes and granular leukocytes begin forming from cytoarchitecturally cognate hematogenetic cells. The cells become distinguishable when hemoglobin starts to accumulate in the erythroblasts and electron dense bodies form in the leukoblasts. Vasculogenesis begins from day 7 in different areas of the developing lung: erthrocytes (but not granular leukocytes) appear to attract committed vasculogenetic cells (angioblasts) that form an endothelial lining and vessel wall. Arrangement of angioblasts around forming blood vessels sets the direction along which the vessels sprout (angiogenesis). In some areas of the developing lung, through what seems like an inductive erythropoietic process, arcades of erythrocytes organize. Once endothelial cells surround such continuities, discrete vascular units organize. By day 10, the major parts of the in-built (intrinsic) pulmonary vasculature are assembled. Complete pulmonary circulation (i.e., through the exchange tissue) is not established until after day 18 when the blood capillaries start to develop. Since the precursory erythrocytes do not have a respiratory role, it is imperative that de novo erythropoiesis is essential for vasculogenesis. Diffuse (fragmentary) development and subsequent piecemeal assembly of the pulmonary vascular system may explicate the fabrication of a complex circulatory architecture that grants cross-current, counter-current, and multicapillary serial arterialization designs in the exchange tissue of the avian lung. The exceptional respiratory efficiency of the avian lung is largely attributable to the geometries (physical interfacing) between the bronchial and vascular elements at different levels of morphological organization.

Animals↗

PubMed2003

Developmental dynamics of the bronchial (airway) and air sac systems of the avian respiratory system from day 3 to day 26 of life: a scanning electron microscopic study of the domestic fowl, Gallus gallus variant domesticus.

The lung buds were first conspicuous on day 3 of embryogenesis. They fused on day 4 and the common growth divided into left and right primordial lungs on day 5. Progressively, the lungs elongated, diverged, and advanced towards the respective dorsolateral aspects of the body wall, reaching their definitive topographical locations in the coelomic cavity on day 6. On day 7, they rotated, attached onto the ribs, gradually started to slide into them, and were deeply inserted by day 8. The primary bronchus (PB) first appeared as a solid cord of epithelial cells (day 4) that successively canalized as it invaded the surrounding mesenchyme, extending along the proximal-distal axis of the lung. From day 8, the secondary bronchi (SB) begun to sprout from the PB in a craniocaudal sequence. On day 9, the parabronchi (PR) started to bud from the SB, projecting into the adjacent mesenchyme. They commenced to canalize on day 10 and greatly increased in length, number, and diameter. By day 13, the PR had anastomosed profusely and totally masked the SB. The luminal surface of the PR was lined by a columnar epithelium from which the atria (day 15), infundibulae (day 16), and air capillaries (ACs) (day 18) developed. At hatching (day 21), the ACs were well developed and had anastomosed profusely with the blood capillaries. Of the air sacs (ASs), the abdominal ones appeared earliest (day 5) followed by the cervical ones on day 6. In quick succession, the other ASs were well formed by day 10. After hatching, no further consequential structures formed: only shifts in topographical locations and an increase in size and number occurred. Morphogenetically, the avian respiratory system differs from the mammalian one in certain key aspects: besides the ASs that are unique to it, the lung is exceptionally complex in structure and is essentially mature at the end of the embryonic life.

Air Sacs↗

PubMed2003

A systematic study of the development of the airway (bronchial) system of the avian lung from days 3 to 26 of embryogenesis: a transmission electron microscopic study on the domestic fowl, Gallus gallus variant domesticus.

In the embryo of the domestic fowl, Gallus gallus variant domesticus, the lung buds become evident on day 3 of development. After fusing on the ventral midline, the single entity divides into left and right primordial lungs that elongate caudally while diverging and shifting towards the dorsolateral aspects of the coelomic cavity. On reaching their definitive topographical locations, the lungs rotate along a longitudinal axis, attach, and begin to slide into the ribs. First appearing as a solid cord of epithelial cells that runs in the proximal-distal axis of the developing lung, progressively, the intrapulmonary primary bronchus begins to canalize. In quick succession, secondary bronchi sprout from it in a craniocaudal sequence and radiate outwards. On reaching the periphery of the lung, parabronchi (tertiary bronchi) bud from the secondary bronchi and project into the surrounding mesenchymal cell mass. The parabronchi canalize, lengthen, increase in diameter, anastomose, and ultimately connect the secondary bronchi. The luminal aspect of the formative parabronchi is initially lined by a composite epithelium of which the peripheral cells attach onto the basement membrane while the apical ones project prominently into the lumen. The epithelium transforms to a simple columnar type in which the cells connect through arm-like extensions and prominently large intercellular spaces form. The atria are conspicuous on day 15, the infundibulae on day 16, and air capillaries on day 18. At hatching (day 21), the air and blood capillaries have anastomosed profusely and the blood-gas barrier become remarkably thin. The lung is well developed and potentially functionally competent at the end of the embryonic life. Thereafter, at least upto day 26, no further consequential structures form. The mechanisms by which the airways in the avian lung develop fundamentally differ from those that occur in the mammalian one. Compared with the blind-ended bronchial system that inaugurates in the mammalian lung, an elaborate, continuous system of air conduits develops in the avian one. Further studies are necessary to underpin the specific molecular factors and genetic processes that direct the morphogenesis of an exceptionally complex and efficient respiratory organ.

Animals↗

PubMed2003

Expression of fibroblast growth factor-2 (FGF-2) in early stages (days 3-11) of the development of the avian lung, Gallus gallus variant domesticus: an immunocytochemical study.

In the avian lung, the bronchial system forms from epithelial (endodermal) cells. The intrapulmonary primary bronchus is the focal point of airway development. It originates secondary bronchi (SB) along its proximal-distal extent and parabronchi (tertiary bronchi) arise from and connect the SB. From as early as day 3.5, fibroblast growth factor-2 (FGF-2) is diffusely expressed in the epithelial and mesenchymal cells. Up-regulation of FGF-2 in discrete areas of the developing lung seem to set the growth rate, trajectories followed, areas appropriated, three-dimensional symmetry and coupling of the airways. Expressed early in development and persisting into the incubation period, FGF-2 may be involved in the formation of the avian lung. Morphogenetic differences between the avian and the mammalian lungs may explain the existing structural contrarieties.

Animals↗

PubMed2002

Fundamental structural aspects and features in the bioengineering of the gas exchangers: comparative perspectives.

Over its life, an organism's survival and success are determined by the inventory of vital adaptations that its progenitors have creatively appropriated, devised and harnessed along the evolutionary pathway. Such conserved attributes provide the armamentarium necessary for withstanding the adverse effects of natural selection. Refinements of the designs of the respiratory organs have been critical for survival and phylogenetic advancement of animal life. Gas exchangers have changed in direct response to the respiratory needs of whole organisms in different environmental states and conditions. Nowhere else is the dictum that in biology 'there are no rules but only necessities' more manifest than in the evolutionary biology of the gas exchangers. The constructions have been continually fashioned and refined to meet specific needs. Solutions to common respiratory needs have been typified by profound structural convergence. Over the evolutionary continuum, as shifts in environmental situations occurred, infinitely many designs should theoretically have emerged. Moreover, without specific selective pressures and preference for certain designs, considering that there are only two naturally occurring respirable fluid media (air and water), air-lungs, water-lungs, air-gills and water-gills would have formed to similar extents. Factors such as body size, phylogenetic level of development, respiratory medium utilized and habitats occupied have permutatively prescribed the design of the gas exchangers. The construction of the modern gas exchangers has eventuated through painstaking cost-benefit analysis. Trade-offs and compromises have decreed only a limited number of structurally feasible and functionally competent outcomes. The morphological congruity (analogy) of the gas exchangers indicates that similar selective pressures have compelled the designs. Solutions to metabolic demands for molecular O2 have only differed in details. Passive physical diffusion, for example, is the ubiquitous method of transfer of O2 across biological tissues. Gills, evaginated gas exchangers, were the primordial respiratory organs that evolved for water breathing, whereas lungs (invaginated gas exchangers) developed for terrestrial (air) breathing. Transitional (= bimodal = amphibious) breathing has evolved in animals with specialized organs that extract O2 from both water and air. Lungs are tidally (= bidirectionally) ventilated, while gills are unidirectionally ventilated, a feature that allows the highly efficient counter-current disposition between blood and water. Since animals occupy inconstant environmental milieus and their metabolic states vary, gas exchangers are designed to operate optimally across a spectrum of conditions that range from resting to exercise and even under hypoxia. Inbuilt structural and functional flexibility provides the requisite safety factors that allow adjustments to modest pressures. The fundamental structural features that determine the respiratory function of a gas exchanger are respiratory surface area, thickness of the blood-water/gas (tissue) barrier and volume of the pulmonary capillary blood. The diffusing capacity of a gas exchanger correlates directly with the surface area and inversely with the thickness of the blood-water/gas (tissue) barrier. An extensive surface area is generated in gills by extensive stratification of the gas exchanger and in lungs by profuse internal subdivision. Compartmentalization yields small terminal gas exchange compartments that compel greater commitment of energy to ventilate. The surfactant, a phospholipid lining, reduces the forces of surface tension at the air-water interface. This attenuates the propensity of physical collapse of the minute gas exchange units and minimizes the cost of ventilation. The surfactant characterizes all the gas exchangers derived from the piscine air bladder. In the lower air-breathing vertebrates, such as the lungfishes (Dipnoi), amphibians and certain reptiles, the pneumocytes are not differentiated into type I and II cells, as is the case in the lungs of the higher vertebrates-birds and mammals. It is envisaged that in endotherms, the overall numerical density of the pneumocytes and hence the O2 consumption of the gas exchangers may be reduced and a thin blood-gas (tissue) barrier generated, factors that enhance respiratory efficiency. The thin blood-gas (tissue) barriers, for example, those of the mammalian (in the respiratory sections of the interalveolar septum) and avian lungs, consist of an epithelial cell and an endothelial cell with a common basement membrane. An interstitial space occurs in the blood-air/water (tissue) barriers of the gas exchangers of fish gills and lungs of lungfishes, amphibians, reptiles and in the supportive parts of the interalveolar septum of the mammalian lung. Collagen, elastic tissue, nerves, lymphatic vessels and smooth muscle elements are found in the interstitial space. The thickness of the blood-air/water (tissue) barrier allometrically changes very little. This suggests that the thicknesses of the blood-water/air (tissue) barriers have been optimized. The presentation and exposure to the gas exchange media (water/air to blood), features dictated by the geometry and arrangements of the structural components of the gas exchangers, contribute greatly to respiratory efficiency. The countercurrent presentation between water and blood in fish gills is the most efficient design in the evolved gas exchangers: It was imperative for survival in water, a medium that contains relatively less O2 and is more expensive to breathe. In the evolved vertebrate gas exchangers, the exposure of blood to air is best manifested in the diffuse design of the avian lung, where the capillary blood is literally suspended in a three-dimensional air space, the blood being exposed to air virtually across the entire blood-gas (tissue) barrier. A double capillary design occurs in the lungs of amphibians and generally those of reptiles, whereas a single capillary design commonly occurs in those of adult mammals. The capillary loading (the ratio of the volume of the capillary blood to the surface area across which blood is exposed to air) in lungs with a double capillary arrangement is high and manifests a poor design. On the other hand, the low capillary loading that characterizes the single capillary system indicates better exposure of blood to air and greater respiratory capacity. Fractal geometry features in the construction of the gas exchangers. The highly versatile design allows the gas exchangers to function optimally under different conditions and circumstances and to maintain congruent morphologies over a wide range of body size, shape and metabolic capacities. At the gas exchange level, sheet-flow design preponderates in the evolved gas exchangers; blood is efficiently exposed to the external respiratory medium. The respiratory capacity of a gas exchanger is comprehensively granted by refinements of structural features and functional processes. Modelling, mathematical integration of structural and functional parameters, provides a holistic view of the essence of the design of a gas exchanger.

Animals↗

PubMed2002

Some recent advances on the study and understanding of the functional design of the avian lung: morphological and morphometric perspectives.

The small highly aerobic avian species have morphometrically superior lungs while the large flightless ones have less well-refined lungs. Two parabronchial systems, i.e. the paleopulmo and neopulmo, occur in the lungs of relatively advanced birds. Although their evolution and development are not clear, understanding their presence is physiologically important particularly since the air- and blood flow patterns in them are different. Geometrically, the bulk air flow in the parabronchial lumen, i.e. in the longitudinal direction, and the flow of deoxygenated blood from the periphery, i.e. in a centripetal direction, are perpendicularly arranged to produce a cross-current relationship. Functionally, the blood capillaries in the avian lung constitute a multicapillary serial arterialization system. The amount of oxygen and carbon dioxide exchanged arises from many modest transactions that occur where air- and blood capillaries interface along the parabronchial lengths, an additive process that greatly enhances the respiratory efficiency. In some species of birds, an epithelial tumescence occurs at the terminal part of the extrapulmonary primary bronchi (EPPB). The swelling narrows the EPPB, conceivably allowing the shunting of inspired air across the openings of the medioventral secondary bronchi, i.e. inspiratory aerodynamic valving. The defence stratagems in the avian lung differ from those of mammals: fewer surface (free) macrophages (SMs) occur, the epithelial cells that line the atria and infundibula are phagocytic, a large population of subepithelial macrophages is present and pulmonary intravascular macrophages exist. This complex defence inventory may explain the paucity of SMs in the avian lung.

Animals↗

PubMed2002

Composite cellular defence stratagem in the avian respiratory system: functional morphology of the free (surface) macrophages and specialized pulmonary epithelia.

Qualitative and quantitative attributes of the free respiratory macrophages (FRMs) of the lung--air sac systems of the domestic fowl (Gallus gallus variant domesticus) and the muscovy duck (Cairina moschata) were compared with those of the alveolar macrophages of the lung of the black rat (Rattus rattus). The birds had significantly fewer FRMs compared to the rat. In the birds, the FRMs were found both in the lungs and in the air sacs. Under similar experimental conditions, the most robust FRMs were those of the domestic fowl followed by those of the rat and the duck. Flux of macrophages onto the respiratory surface from the subepithelial compartment and probably also from the pulmonary vasculature was observed in the birds but not in the rat. In the duck and the domestic fowl, a phagocytic epithelium that constituted over 70% of the surface area of the blood-gas (tissue) barrier lines the atrial muscles, the atria and the infundibulae. The epithelial cells of the upper respiratory airways contain abundant lysosomes, suggesting a high lytic capacity. By inference, the various defence strategies in the avian lung may explain the dearth of FRMs on the respiratory surface. We counter-propose that rather than arising directly from paucity of FRMs, an aspect that has been over-stressed by most investigators, the purported high susceptibility of birds (particularly table birds) to respiratory ailments and afflictions may be explained by factors such as inadequate management and husbandry practices and severe genetic manipulation for fast growth and high productivity, manipulations that may have weakened cellular and immunological defences.

Air Sacs↗

PubMed2002

Structure, function and evolution of the gas exchangers: comparative perspectives.

Over the evolutionary continuum, animals have faced similar fundamental challenges of acquiring molecular oxygen for aerobic metabolism. Under limitations and constraints imposed by factors such as phylogeny, behaviour, body size and environment, they have responded differently in founding optimal respiratory structures. A quintessence of the aphorism that 'necessity is the mother of invention', gas exchangers have been inaugurated through stiff cost-benefit analyses that have evoked transaction of trade-offs and compromises. Cogent structural-functional correlations occur in constructions of gas exchangers: within and between taxa, morphological complexity and respiratory efficiency increase with metabolic capacities and oxygen needs. Highly active, small endotherms have relatively better-refined gas exchangers compared with large, inactive ectotherms. Respiratory structures have developed from the plain cell membrane of the primeval prokaryotic unicells to complex multifunctional ones of the modern Metazoa. Regarding the respiratory medium used to extract oxygen from, animal life has had only two choices--water or air--within the biological range of temperature and pressure the only naturally occurring respirable fluids. In rarer cases, certain animals have adapted to using both media. Gills (evaginated gas exchangers) are the primordial respiratory organs: they are the archetypal water breathing organs. Lungs (invaginated gas exchangers) are the model air breathing organs. Bimodal (transitional) breathers occupy the water-air interface. Presentation and exposure of external (water/air) and internal (haemolymph/blood) respiratory media, features determined by geometric arrangement of the conduits, are important features for gas exchange efficiency: counter-current, cross-current, uniform pool and infinite pool designs have variably developed.

Adaptation, Physiological↗

PubMed2001

Morphometric characterization of the airway and vascular systems of the lung of the domestic pig, Sus scrofa: comparison of the airway, arterial and venous systems.

The bronchial system (BS), the pulmonary artery (PA) and the pulmonary vein (PV) of the lung of the domestic pig, Sus scrofa were simultaneously cast with silicone rubber and studied. Asymmetrical dichotomous bifurcation preponderated in the tree-like arrangement of the three conducting systems. Lengths and diameters of the various generations were measured. At the extremities of the BS and the PA, alveoli and blood capillaries related very closely. In the cranial and middle lobes of the right and left lungs, topographically, the PA and the PV closely followed the BS, but in the accessory and the caudal (diaphragmatic) lobes, only the PA accompanied the BS: the PV run intersegmentally. Certain similarities and differences were observed between the diameters and lengths of the various generations of the three conducting systems. The strong correlations between some of the structural parameters indicated a high level of structural optimization. While morphometric variations suggest that the air and the blood flow dynamics may somewhat differ between the three conducting systems, they may also register structural features unique to the lung of the domestic pig, an animal that has been highly genetically exploited for fast growth and now leads an indolent lifestyle in captivity.

Animals↗

PubMed2001

A qualitative and quantitative study of the lung of an ostrich, Struthio camelus.

The ostrich lung, with its lack of interparabronchial septa, the presence of very shallow atria and exceptional morphometric refinement, structurally resembles those of small, energetic flying birds, whereas it also displays features characteristic of the flightless ratites in which the neopulmo is relatively poorly developed and a segmentum accelerans may be generally lacking. The large size of the bronchial system of the ostrich may help explain the unique shifts in the airflow pathways that must occur from resting to panting breathing, explaining its insensitivity to acid-base imbalance of the blood during sustained panting under thermal stress. The mass-specific volume of the lung is 39.1 cm(3)kg(-1) and the volume density of the exchange tissue is remarkably high (78.31%). The blood-gas (tissue) barrier is relatively thick (0.56 microm) but the plasma layer is very thin (0.14 microm). In this flightless ratite bird, the mass-specific surface area of the tissue barrier (30.1 cm(2)g(-1)), the mass-specific anatomical diffusing capacity of the tissue barrier for oxygen (0.00 22 ml O(2) s(-1) Pa(-1) kg(-1)), the mass-specific volume of pulmonary capillary blood (6.25 cm(3)kg(-1)) and the mass-specific total anatomical diffusing capacity for oxygen (0.00073 ml O(2) s(-1) Pa(-1) kg(-1)) are equivalent to or exceed those of much smaller highly aerobic volant birds. The distinctive morphological and morphometric features that seem to occur in the ostrich lung may explain how it achieves and maintains high aerobic capacities and endures long thermal panting without experiencing respiratory alkalosis.

Animals↗

PubMed2000

Comparative respiratory morphology: themes and principles in the design and construction of the gas exchangers.

Along the evolutionary continuum, a kaleidoscope of gas exchangers has evolved from the simple cell membrane of the primeval unicells. The most momentous events in this process were: the intensification of molecular oxygen in the biosphere and its appropriation into aerobic metabolism, the rise of multicellular organisms, the development of a circulatory system and carrier pigments in blood, the advocacy of air breathing, adoption of suctional breathing, and the shift to endothermy. To satisfy species-specific needs for oxygen, some constraints were overcome through transactions that obliged certain compromises and trade-offs. Optimal designs of the gas exchangers for particular phylogenetic levels of development, habitat, and lifestyle have developed only so far as to satisfy prescribed needs. The efficiency of the human lung, for example, falls well below those of certain taxa that are considered to be relatively "less advanced." Utilizing different resources and strategies, in fascinating processes of conformity, different groups of animals have developed similar respiratory structures. In most cases, the analogy reflects evolutionary convergence in response to corresponding selective pressures rather than common ancestry. Anat Rec (New Anat) 261:25-44, 2000.

Animals↗

PubMed2000

Is the sheet-flow design a 'frozen core' (a Bauplan) of the gas exchangers? Comparative functional morphology of the respiratory microvascular systems: illustration of the geometry and rationalization of the fractal properties.

The sheet-flow design is ubiquitous in the respiratory microvascular systems of the modern gas exchangers. The blood percolates through a maze of narrow microvascular channels spreading out into a thin film, a "sheet". The design has been convergently conceived through remarkably different evolutionary strategies. Endothelial cells, e.g. connect parallel epithelial cells in the fish gills and reptilian lungs; epithelial cells divide the gill filaments in the crustacean gills, the amphibian lungs, and vascular channels on the lung of pneumonate gastropods; connective tissue elements weave between the blood capillaries of the mammalian lungs; and in birds, the blood capillaries attach directly and in some areas connect by short extensions of the epithelial cells. In the gills, skin, and most lungs, the blood in the capillary meshwork geometrically lies parallel to the respiratory surface. In the avian lung, where the blood capillaries anastomose intensely and interdigitate closely with the air capillaries, the blood occasions a 'volume' rather than a 'sheet.' The sheet-flow design and the intrinsic fractal properties of the respiratory microvascular systems have produced a highly tractable low-pressure low-resistance region that facilitates optimal perfusion. In complex animals, the sheet-flow design is a prescriptive evolutionary construction for efficient gas exchange by diffusion. The design facilitates the internal and external respiratory media to be exposed to each other over an extensive surface area across a thin tissue barrier. This comprehensive design is a classic paradigm of evolutionary convergence motivated by common enterprise to develop corresponding functionally efficient structures. With appropriate corrections for any relevant intertaxa differences, use of similar morphofunctional models in determining the diffusing capacities of various gas exchangers is warranted.

Animals↗

PubMed2000

Functional morphology of the gas-gland cells of the air-bladder of Oreochromis alcalicus grahami (teleostei: cichlidae): an ultrastructural study on a fish adapted to a severe, highly alkaline environment.

Oreochromis alcalicus grahami is a small cichlid fish that lives in shallow peripheral lagoons of Lake Magadi, Kenya. The internal surface of the air-bladder is highly vascularized, illustrating possible utilization as an accessory respiratory organ. The wall of the bladder consists of five distinct tissue layers. From the outer to the inner surfaces are: a squamous, undifferentiated epithelial cell; a collagen-elastic tissue space; a smooth muscle tissue band; an inner connective tissue space; and columnar gas-gland cells projecting into the lumen. The cell membrane over the perikarya of the gas-gland cells was sporadically broken. The disruptions were ascribed to possible physical damage by discharging gas-bubbles. Juxtaluminally, the gas-gland cells attached across tight junctions. The cells have highly euchromatic nuclei and conspicuously large intracytoplasmic secretory bodies. On the blood capillary facing (basal) aspect, the cell membrane of the gas-gland cells is highly amplified. The cells insert onto a smooth muscle tissue band. The morphological features and the topographical arrangement of the gas-gland cells in O. a. grahami are indicative of an operative exchange of materials between them and the underlying tissue components especially the blood capillaries. For a fish that subsists in hot, highly saline and alkaline water heavily invested by avian predators and where the partial pressure of oxygen diurnally shifts from virtual anoxia to hyperoxia, development of a versatile air-bladder for efficient buoyancy control conforms to the functional demands placed on it by a unique environment. Illustratively, instead of the gas-gland morphology in O. a. grahami resembling that in the freshwater fishes, the group from which the fish has evolved, it compares more closely to that of the marine fish. This similarity suggests amazing parallel evolutionary adaptation to biophysically corresponding aquatic milieus.

Africa↗

PubMed2000

Inspiratory aerodynamic valving in the avian lung: functional morphology of the extrapulmonary primary bronchus.

The form, geometry and epithelial morphology of the extrapulmonary primary bronchi (EPPB) of the domestic fowl (Gallus gallus var. domesticus) and the rock dove (Columba livia) were studied microscopically and by three-dimensional computer reconstruction to determine the structural features that may be involved in the rectification of the inspired air past the openings of the medioventral secondary bronchi (MVSB), i.e. the inspiratory aerodynamic valving (IAV). In both species, the EPPB were intercalated between the clavicular and the cranial thoracic air-sacs. A notable difference between the morphology of the EPPB in G. g. domesticus and C. livia was that, in the former, the EPPB were constricted at the origin of the MVSB, while a dilatation occurred at the same site in the latter. In both species, a highly vascularized, dorsally located hemispherical epithelial swelling was observed cranial to the origin of the MVSB. The MVSB were narrow at their origin and variably angled relative to the longitudinal axis of the EPPB. Conspicuous epithelial tracts and folds were observed on the luminal aspect of the EPPB in both C. livia and G. g. domesticus. From their marked development and their orientation relative to the angled MVSB, these properties may influence the flow of the air in the EPPB. It was concluded that features such as syringeal constriction, an intimate topographic relationship between the EPPB and the cranial air-sacs, prominent epithelial tracts and folds, an epithelial swelling ahead of the origin of the first MVSB (corresponding to the 'segmentun accelerans'), and narrowing and angulation of the MVSB at their origin, may together contribute to IAV to a variable extent. In as much as the mechanism of pulmonary ventilation and mode of airflow in the parabronchial lung are basically similar in all birds, the morphological differences observed between G. g. domesticus and C. livia suggest that either the mechanism of production of IAV or its functional efficiency may be different, at least in these two species of birds.

Animals↗

PubMed2000

What it takes to fly: the structural and functional respiratory refinements in birds and bats.

In absolute terms, flight is a highly energetically expensive form of locomotion. However, with respect to its cost per unit distance covered, powered flight is a very efficient mode of transport. Birds and bats are the only extant vertebrate taxa that have achieved flight. Phylogenetically different, they independently accomplished this elite mode of locomotion by employing diverse adaptive schemes and strategies. Integration of functional and structural parameters, a transaction that resulted in certain trade-offs and compromises, was used to overcome exacting constraints. Unique morphological, physiological and biochemical properties were initiated and refined to enhance the uptake, transfer and utilization of oxygen for high aerobic capacities. In bats, exquisite pulmonary structural parameters were combined with optimal haematological ones: a thin blood-gas barrier, a large pulmonary capillary blood volume and a remarkably extensive alveolar surface area in certain species developed in a remarkably large lung. These factors were augmented by, for example, exceptionally high venous haematocrits and haemoglobin concentrations. In birds, a particularly large respiratory surface area and a remarkably thin blood-gas (tissue) barrier developed in a small, rigid lung; a highly efficient cross-current system was fabricated within the parabronchi. The development of flight in only four animal taxa (among all the animal groups that have ever evolved; i.e. insects, the now-extinct pterosaurs, birds and bats) provides evidence for the enormous biophysical and energetic constraints that have stymied volancy. Bats improved a fundamentally mammalian lung to procure the large amounts of oxygen needed for flight. The lung/air sac system of birds is not therefore a prescriptive morphology for flight: the essence of its design can be found in the evolution of the reptilian lung, the immediate progenitor stock from which birds arose. The attainment of flight is a classic paradigm of the remarkable adaptability inherent in organismal and organic biology for countering selective pressures by initiating elegant morphologies and physiologies.

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