The new adapatational biology mainly germinated spontaneously from several marginal areas of ecology, instead of from marginal areas of physiology or molecular biology, and this has a special reason. The new direction is based on the combination of Darwinian paradigm and microbiology, and it needs a nonrestricted space ready for the integration of knowledge and theory. Ecology is in a special position and can play a special role. On the one hand, ecology naturally mingles with microbiology. On the other hand, it has an innate theoretical connection with the evolution. In consequence, ecology is a fertile and special soil on which many borderland disciplines germinate and grow, such as the life-history evolutionary ecology, ecology of behavioral strategy, physioecology of stress resistance, and physioecology of phenotypic plasticity. Although evolutionary pathophysiology originates from physiology, during the process associating with evolutionary biology, it unconsciously goes to a new study of disease's physiological and ecological causations.
Among these branches, the study of bionomic strategy bears the most influence, and its creation represents a typical history breaking away from the modern synthesis. The idea of bionomic strategy was not deliberately coined to comply with the principle of intraspecies comparison. Its origin can be traced back to an important debate on group selection or individual selection. George Christopher Williams, an American evolutionary biologist
, insisted that the unit of natural selection is individual. He published a book called Adaptation and Natural Selection
in the early 1960s with a vigorous critique of group selection
. His main point is that a trait with possible long-term benefit to the species or groups, but useless and even harmful to the individual, could not have selective advantage. The debate generated a great effect, and people may notice that the individual selection, which is opposite to the group selection, is the foundation of modern selfish-gene theory. From this point of view, Williams' work appears to be conducive to the sexual selection determinism and supports the position of the modern synthesis.
However, it is often neglected that G. C. Williams also attached great importance to phenotypic physiology. As a marine ecologist, he was the first scholar who put forward the aging-related and adaptive evolutionism in 1957, and after that he further deepened his thinking about individual adaptation. He noted that the definition of adaptation in the current theories of evolution was ambiguous, and a more explicit explanation and understanding for the concept of adaptation was important. The contemporary view of evolutionary adaptation is just a simple and naive prospect, and its defects need to be modified with a more explicit theory. The research on evolution needs a theory about adaptation. He argued that quantitative research on the adaptive phenomena was required. Rather than studying the superficial ecad adaptation in traditional taxonomy, he suggested that we should study the employment of general strategies in life process and the variance in phylogenesis. In particular, he mentioned the variations in life history phylogenesis, such as paedogenesis, parthenogenesis, apomixis, the alternation of generations, abnormal animals etc. G. C. Williams might be the first person who was sensitive to the new issues about adaptive physiological problems.
Coincidentally, the controversy over individual selection has drawn the attention of another scholar, Maynard Smith. Smith was proficient in mathematics and built a mathematical model for individual selection. Smith was the person who first introduced the game theory to explain evolutionism. His quantitative calculation was initially aimed at explaining why the essentially selfish individuals were able to coexist in peace and even cooperate with each other. His conclusion of the calculations is still controversial, but the calculation has opened up the door for research combining the concept of strategy with the comparative study. Smith himself was not aware of the historic significance of his theory at that time. In consequence, the comparative study on bionomic strategy developed rapidly, focusing on the selection of phenotypic adaptive variations and life process variations which were advocated by G. C. Williams. This is definitely beyond the expectation of Maynard Smith, who had made a historic contribution. This sort of unexpected result is not uncommon in the history of science.
At the beginning of the study of bionomic strategy, the task chiefly included comparing differences in interspecies or sibling species. Interspecies adaptation comparison is the normal paradigm in traditional evolutionary biology, and it was especially created from the issue of individual vs. group selection. Therefore, the study of bionomic strategy was primarily regarded as a complement or component of the modern synthesis. Many research results and calculations have been published in this aspect, and some important theoretical thinking has been put forward, such as the r/K selection theory. However, the study of bionomic strategy was soon naturally integrated into the intraspecies comparative system. Because this paradigm is very suitable and convenient to be used to explain the variations in life history phylogenesis which G. C. Williams was concerned about, such as the adaptive values of paedogenesis, parthenogenesis, apomixis, the alternation of generations, and the abnormal animals. In other words, the theoretical analytic method for exploring the physiological adaptive variation in individual phenotypes actually comes into being, living up to G. C. Williams' expectation.
Then a method of intraspecific adaptation comparison was established by observing bionomic strategy. The method was at first successfully introduced to the comparative study of intraspecies life history, then further extended to searching the adaptive value of intraspecific behavioral variations. A great deal of behavioral strategy studies in behavioral ecology are part of this category, such as the study of foraging strategy, fighting strategy, territory behavioral strategy, etc.
The most important significance of the strategy studies is that it introduces the Darwinian paradigm into the investigation on issues of intraspecific adaptive variations in phenotype and physiology. The study of bionomic strategy inherited the traditional Darwinian methodology using the new concept of bionomic strategy as a comparative framework, and successfully employed the mathematical model to meet the needs of explaining intraspecies comparison. That is, using small hypothesis to set a model for the given objects, quantifying traits of the objects as much as possible to a series of variable parameters, calculating them to make a prediction, then comparing the closeness of the actual observations with the calculating results of the model. The term strategy breaks the boundary between physiology and ecology, being a combined expression for both. Its appearance provides a suitable concept for the explanation of intraspecific adaptive variation in physiological systems, that is, strategy is the basal object in analysis of physiological adaptive variation.
At present, the major branches of adaptational biology have different backgrounds, and each of them has its own characteristics in concept employment and theoretical analytical method. Until now an incorporated systematic framework for them has not formed. However, all of them have coincidentally developed a way of thinking similar to the bionomic strategy, such as the concept stress resistance in stress biology, compensation and defense in evolutionary pathophysiology, and so on. Therefore, we can consider them as a combination of a broader strategy concept with microbiology knowledge, which viewed from the perspective of molecule-level and biochemistry-level can explain and calculate the adaptive value of an individual organism's physiological variations under different environmental conditions. This way of thinking expanded rapidly and covered a wide range. Although a unified theoretical paradigm has not yet been formed, the concept of strategy has such theoretically cover-all ability that the branches have a tendency to form a common and broader concept of strategy. At present, this process is still underway.