Summary and Conclusion

Abstract

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 ex- changers 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 task of designing or developing an efficient structure is not an easy one. In biology, millions of years of natural selection have provided an encyclopedic variety of imperfect or intermediate structures. Gordon (1988)