On adaptation, the assessment of adaptations, and the value of adaptive arguments in phylogenetic reconstruction

Evolutionary adaptation concerns a relative concept and the study of adaptations is directed to structures of individuals. The concept is devoid of any meaning when it is applied to species or populations. Adaptation is not synonymous with fitness or survival but does contribute to both of them. The term adaptation has a dual meaning since it refers both to the process of adaptation and to the state of being adapted. In the process of adaptation the mechanism of natural selection takes a prominent position. But the operation and effectiveness of natural selection are constrained by various limiting factors. Besides that, features may also be the result of nonadaptive evolution and only attain their present adaptive function at a later point in time. Another possibility is that features have at present a function different from the one for which they were initially designed. With respect to the state of being, the study of adaptation attempts to examine whether a particular feature indeed forms an adequate response to selection forces from the environment. Five methods or approaches generally are used to assess the adaptive significance of features, viz. the comparative, correlation, optimization, cladistic, and synthetic approach. Only the last-mentioned approach forms an adequate method since it attempts to establish, by direct analysis, which well-defined selection force exerts its influence on a certain character. The practicing taxonomist is faced with the problem that the data necessary to apply the synthetic method, generally require detailed field studies. Not all evolutionary changes are under the influence of natural selection. The presence of some features may be based on entirely different mechanisms, such as genetic drift, mutational pressure, pleiotropic gene action, allometric growth, or ecophenotypic responses. Various problems inherent to the optimization approach, and several others of practical and theoretical nature, make the morphocline method of the functional and evolutionary morphologists unsuitable as a method for phylogenetic reconstruction.     

BIOCHEMICAL ADAPTATION AND LOSS OF GENETIC CAPACITY IN HELMINTH PARASITES

1. Adaptation and loss of genetic capacity differ chiefly in that adaptation is goal- directed whereas loss of genetic capacity is not. Given sufficient information about an individual organism and its environment, adaptations are recognizable without reference to historical events extending beyond a single generation. This is not true of loss of genetic capacity, which requires a preliminary judgement that genetic information now absent was present in ancestral organisms. Together, adaptation and loss of genetic capacity are the major contributors to overall reproductive fitness. Accidental selection is genetically associated with adaptation, but is not goal-directed. 2. Adaptations arevariant or invariant; invariant adaptations comprising biochemical unity, and variant adaptations contributing to biochemical diversity. Variant adaptations may be either exploitive or epigenetic. Exploitive adaptations are a measure of thegenetic capacity for phenotypic response to an altered environment, which the individual may not in fact encounter. Epigenetic adaptations are more rigidly programmed and are responsive to altered environments only insofar as these are a constant feature of the life cycle. 3. Selected observations in the biochemistry of helminth parasites are examined with respect to their interpretation in terms of adaptation, loss of genetic capacity and accidental selection. Secure judgements concerning adaptation are often possible at the most general level, i.e. when the physicochemical properties of the environment, such as temperature or oxygen supply, are clearly defined. I t is more difficult to make judgements concerning the specific mechanisms used in achieving these goals. Conclusions concerning loss of genetic capacity require knowledge of the specific function through-out the life cycle. In many cases loss of genetic capacity is only apparent, as the function appears in another part of the life cycle. Such apparent losses are in reality epigenetic adaptations. These concepts are helpful in interpreting past work and in devising new experiments. 4. Development in helminth parasites includes a pronounced capacity for the orderly release of information to be used in the next stage. As each stage may require a radically different environment, programming for it may lead to phenomena which are superficially puzzling, such as the existence of aerobic electron transport systems in a stage whose energy metabolism is fermentative. The concept of epigenetic adaptation is especially useful for interpreting such observations. 5. Although possible adaptations are most readily apparent in biochemically complex mechanisms, these mechanisms are an expression of the orderly effects of many different primary gene products which have not been much studied. There are indications that organisms possessing relatively complex life cycles may provide opportunities for relating primary gene products, such as isozymes, to their physiological functions.     

Saltatory Processes and Altricial to Precocial Forms in the Ontogeny of Fishes

SYNOPSIS. Development is not a gradual but a saltatory process.A combination of qualitative changes in form and function—thresholds—createsboundaries between a succession of quantitative intervals—steps.Thresholds can be modified by an altered time of appearanceof structures and functions (heterochrony), especially duringearly ontogeny, to form an operational basis for the prolongationof juvenile characters and adaptability into later ontogeny.Whereas such prolongation enables juvenilization in phylogeny,analogous principles may operate on a much shorter time scaleto produce the r-selection-like altricial and the K-selection-likeprecocial trends in ontogeny. The inherited capacity to adjustconstantly to the environment (heterochrony) selects for structural,biochemical and behavioral improvements. The tendency is towardthe precocial but the way back to altricial forms is left open.The heterochronous adaptations in early ontogeny can "reverse"the vulnerable specialization, should the environment becomeless stable and/or the community less competitive. Juvenilization,capable of turning gerontomorphosis into paedomorphosis in evolution,and heterochronous shifts of character anlagen, capable of turninga precocial trend into an altricial trend in ontogeny are bothpart of the same biological process which operates during earlyontogeny.     

Capturing the superorganism: a formal theory of group adaptation

Adaptation is conventionally regarded as occurring at the level of the individual organism. However, in recent years there has been a revival of interest in the possibility for group adaptations and superorganisms. Here, we provide the first formal theory of group adaptation. In particular: (1) we clarify the distinction between group selection and group adaptation, framing the former in terms of gene frequency change and the latter in terms of optimization; (2) we capture the superorganism in the form of a ‘group as maximizing agent’ analogy that links an optimization program to a model of a group‐structured population; (3) we demonstrate that between‐group selection can lead to group adaptation, but only in rather special circumstances; (4) we provide formal support for the view that between‐group selection is the best definition for ‘group selection’; and (5) we reveal that mechanisms of conflict resolution such as policing cannot be regarded as group adaptations.     

Information Processing by Cyanobacteria during Adaptation to Environmental Phosphate Fluctuations

Phosphate limited grown Anabaena variabilis has the capability of processing information about external phosphate fluctuations by means of interconnected adaptive events. Adaptive events are physiological processes that are characterized by two opposite manifestations, namely adapted states and adaptive operation modes. In adapted states the energy-converting constituents of the uptake system operate under the prevailing external conditions in a coherent manner with least energy dissipation. Adaptive operation modes take place when adapted states are disturbed by persistent changes in phosphate supply. In this mode the outcome of former adaptations to elevated phosphate levels guides the emergence of a new adapted state. The influence of antecedent adapted states on subsequent adaptations was studied experimentally and characteristic examples for such information processing are given. The theory of self-referential systems allowed analyzing these examples. For this purpose adaptive events had to be considered as elements of a communicating network, in which, along a historic succession of alternating adapted states and adaptive operation modes, information pertaining to the self-preservation of the organism is transferred from one adaptive event to the next: the latter “interprets” environmental changes by means of distinct adaptive operation modes, aimed at preservation of the organism. The result of this interpretation is again leading to a coherent state that is passed on to subsequent adaptive events. A generalization of this idea to the adaptive interplay of other energy converting subsystems of the cell leads to the dynamic view of cellular information processing in which the organism recreates itself in every new experience.Key Words: adaptation, cyanobacteria, information processing, phosphate uptake, self-referential systems     

Semiotics of a Superorganism

Darwinian evolution, as it was first conceived, has two dimensions: adaptation, that is, selection based upon “apt function”, defined as the “good fit” between an organism’s metabolic and biological demands and the environment in which it is embedded; and heredity, the transmissible memory of past apt function. Modern Darwinism has come to focus almost exclusively on hereditary memory, eclipsing the—arguably still-problematic—phenomenon of adaptation. As a result, modern Darwinism retains, at its core, certain incoherencies that, as long as they remain unresolved, preclude the emergence of a fully-coherent theory of evolution. Resolving the incoherencies will involve clarifying the relationship between embodied memory and apt function. In short, adaptation is a problem of semiotics: the organism must interpret the environment to fit well into it. This is well-illustrated by the constructed environments built by colonies of social insects, such as hives or nests, and the ancillary structures that contain them, forming an organism-like system known as a superorganism. The superorganism is marked by a kind of extended physiology, in that these constructed environments often serve as adaptive interfaces between the nest and ambient environment, and are constructed to manage the matter and energy flows between environments that constitute the process of adaptation. These constructed environments are also semiotic phenomena: interpretive structures, governed by information flow between the member insects and the structures they build. I review our findings on one such example: the mounds built by the fungus-cultivating termites of the genus Macrotermes. These structures are dynamic forms that are sustained by flows of soil from deep horizons up into the mound. The form, and hence the function, of the mound is determined by several environmental cues, most notably water and wind, as well as how termites interpret these cues, and signals that flow between termites, both directly and vicariously through the structures they build.     

Mountain birch under multiple stressors – heavy metal‐resistant populations co‐resistant to biotic stress but maladapted to abiotic stress

Stress adaptations often include a trade‐off of weakened performance in nonlocal conditions, resulting in divergent selection, and potentially, genetic differentiation and evolutionary adaptation. Results of a two‐phase (greenhouse and field) common garden experiment demonstrated adaptation of mountain birch (Betula pubescens subsp. czerepanovii) populations from industrially polluted areas of the Kola Peninsula, north‐western Russia, to heavy metals (HM), whereas no adaptations to wind or drought stress were detected in populations from wind‐exposed sites. HM‐adapted seedlings were maladapted to drought but less palatable (co‐resistant) to insect herbivores, even under background HM concentrations. The absence of adaptations to harsh microclimate and the generally high adaptive potential of mountain birch, a critical forest forming tree in subarctic Europe, need to be accounted for in models predicting consequences of human‐driven environmental changes, including the projected climate change.     

Experimental Evolution and Its Role in Evolutionary Physiology

Four general approaches to the study of evolutionary physiology—phylogenetically-basedcomparisons, genetic analyses and manipulations, phenotypicplasticity and manipulation, and selection studies—areoutlined and discussed. We provide an example of the latter,the application of laboratory selection experiments to the studyof a general issue in environmental adaptation, differencesin adaptive patterns of generalists and specialists. A cloneof the bacterium Escherichia coli that had evolved in a constantenvironment of 37°C was replicated into 6 populations andallowed to reproduce for 2,000 generations in a variable thermalenvironment alternating between 32 and 42°C. As predictedby theory, fitness and efficiency of resource use increasedin this new environment, as did stress resistance. Contraryto predictions, however, fitness and efficiency in the constantancestral environment of 37°C did not decrease, nor didthermal niche breadth or phenotypic plasticity increase. Selectionexperiments can thus provide a valuable approach to testinghypotheses and assumptions about the evolution of functionalcharacters.     

Codon Usage in Plastid Genes Is Correlated with Context, Position Within the Gene, and Amino Acid Content

Highly expressed plastid genes display codon adaptation, which is defined as a bias toward a set of codons which are complementary to abundant tRNAs. This type of adaptation is similar to what is observed in highly expressed Escherichia coli genes and is probably the result of selection to increase translation efficiency. In the current work, the codon adaptation of plastid genes is studied with regard to three specific features that have been observed in E. coli and which may influence translation efficiency. These features are (1) a relatively low codon adaptation at the 5′ end of highly expressed genes, (2) an influence of neighboring codons on codon usage at a particular site (codon context), and (3) a correlation between the level of codon adaptation of a gene and its amino acid content. All three features are found in plastid genes. First, highly expressed plastid genes have a noticeable decrease in codon adaptation over the first 10–20 codons. Second, for the twofold degenerate NNY codon groups, highly expressed genes have an overall bias toward the NNC codon, but this is not observed when the 3′ neighboring base is a G. At these sites highly expressed genes are biased toward NNT instead of NNC. Third, plastid genes that have higher codon adaptations also tend to have an increased usage of amino acids with a high G + C content at the first two codon positions and GNN codons in particular. The correlation between codon adaptation and amino acid content exists separately for both cytosolic and membrane proteins and is not related to any obvious functional property. It is suggested that at certain sites selection discriminates between nonsynonymous codons based on translational, not functional, differences, with the result that the amino acid sequence of highly expressed proteins is partially influenced by selection for increased translation efficiency. Received: 21 July 1999 / Accepted: 5 November 1999     

Species of thought: A comment on evolutionary epistemology

The primary outcome of natural selection is adaptation to an environment. The primary concern of epistemology is the acquistion of knowledge. Evolutionary epistemology must therefore draw a fundamental connection between adaptation and knowledge. Existing frameworks in evolutionary epistemology do this in two ways; (a) by treating adaptation as a form of knowledge, and (b) by treating the ability to acquire knowledge as a biologically evolved adaptation. I criticize both frameworks for failing to appreciate that mental representations can motivate behaviors that are adaptive in the real world without themselves directly corresponding to the real world. I suggest a third framework in which mental representations are to reality as species are to ecosystems. This is a many-to-one relationship that predicts a diversity of adaptive representations in the minds of interacting people. As “species of thought”, mental representations share a number of properties with biological species, including isolating mechanisms that prevent them from blending with other representations. Species of thought also are amenable to the empirical methods that evolutionists use to study adaptation in biological species. Empirical studies of mental representations in everyday life might even be necessary for science to succeed as a normative “truth-seeking” discipline.     

Adaptation or exaptation? The case of the human hand

A controversy of relevance to the study of biological form involves the concept of adaptation. This controversy is illustrated by the structure and function of the human hand. A review of the principal definitions of adaptation points to two main problems: (1) they are qualitative and make reference to the whole structure (or substructural feature) and (2) they are based on the idea of natural selection as a moulding factor. The first problem would be solved by a definition that encompasses quantitative measures of the effects of selection, drawing on new advances in the comparative method. The second problem is deeper and presents greater conceptual difficulties. I will argue that the idea of natural selection as a moulding factor depends on the notion of a genetic program for development. But regarding the hand, experimental evidence on limb development challenges the idea of a genetic program for skeletal pattern formation, undermining a simple application of standard adaptationist concepts. These considerations lead to a revised definition of adaptation and interpretation of the evolutionary determinants of the hand’s form.     

Adaptation as process: the future of Darwinism and the legacy of Theodosius Dobzhansky

Conceptions of adaptation have varied in the history of genetic Darwinism depending on whether what is taken to be focal is the process of adaptation, adapted states of populations, or discrete adaptations in individual organisms. I argue that Theodosius Dobzhansky's view of adaptation as a dynamical process contrasts with so-called "adaptationist" views of natural selection figured as "design-without-a-designer" of relatively discrete, enumerable adaptations. Correlated with these respectively process and product oriented approaches to adaptive natural selection are divergent pictures of organisms themselves as developmental wholes or as "bundles" of adaptations. While even process versions of genetical Darwinism are insufficiently sensitive to the fact much of the variation on which adaptive selection works consists of changes in the timing, rate, or location of ontogenetic events, I argue that articulations of the Modern Synthesis influenced by Dobzhansky are more easily reconciled with the recent shift to evolutionary developmentalism than are versions that make discrete adaptations central.     

Prolegomenon for a hypothesis on music as expression of an evolutionary early homeostatic feedback-mechanism. A biomusicological proposal.

Musical experience and creativity are regarded to be largely depending on cultural conditions and hence on higher cognitive functions. True as this may be, there are, however, numerous responses to music-the urge to make music taken into account-which derive from deeper levels of the human organism, namely from arousing alternatively moderating vegetative and limbic functions. Although the behavioral intensity and quality emanating from such evolutionary early nervous structures may be affected by cultural influence, they still seem to be essentially independent. Similar specific responses to acoustical and/or motor patterned stimuli are found among some other higher vertebrates which like humans are equipped with sophisticated mechanisms for hearing, sound production and locomotion, well tuned to each other. However, it is even today an open question whether these manifestations of auditive-phonatory-locomotor abilities just are analogues or if they share a common evolutionary background. The current discussion on this matter has accumulated data which apparently support the latter view pointing to that sexual selection would be the common force, first suggested by Charles Darwin (153). Other, and still more recent data in genetics and neuroscience, may be interpreted as hints at that the common origin would be a more elemental organismic feature, a metabolic-homeostatic variable which due to its evolutionary strength eventually created the platform for a radiation of adaptations concerning species-specific patterned sounds and locomotions with a broad spectrum of tasks, among them sexual selection. This line of reasoning is here, under reference to recent biological data, made the basis for a hypothetical model of music as an expression of an early homeostatic feedback mechanism. Accordingly, in music there is a central variable, a “heart” or a “core”, which is not to be found exclusively in music but appears globally as a releasing mechanism for basic endocrine, autonomous and elementally cognitive functions. It is of acoustical or motoric nature, or of a combination of these characters, and is performed in repetitive trains of impulses. It is further assumed that the target of its operations is mainly proteins with a regulatory effect on the cellular and synaptic states. The principal representatives for these proteins are growth factors, especially the NGF which originally was regarded as a growth stimulator within the peripheral and sympathetical systems but which eventually appeared to be also a synergetic modulator of neuro-endocrine-immuno-reactions, i.e. of the three central homeostatic systems (5, 80). One can speculate that this variable is functionally active at an elemental level such that it has escaped to be knocked out by forceful “higher” and evolutionary younger factors (49:13). This hypothesis — that music has its roots within and is a part of a globally occurring natural acoustical-motoric stimulus, manifested in a great variety of auditory and motoric behavior in humans and among some other higher vertebrates — implies that humankind has developed this stimulus into a category of acoustical structures which oscillate round an instable point of equilibrium. Exactly such structures, not stochastic but neither too predictable, affect the organism mainly on a sensory-vegetative level (59, 102, 137, 151). They are in addition perceptionally optimal in creating cortical space-temporal neural patterns with strong interhemispheric coherence (110, 130). According to this scenario, music did not originate from a human need of communication or as an aspect of sexual selection. It emerged from elemental processes within the individual organism with the aspiration to maintain his bodily and mental fitness, thus on a pre-social level. What was beneficial to the single individual in his fight for survival, was good also for the group and its survival. Starting from that platform music has evolved in symbiosis with dance and play within a large spectrum of social functions, where sexual selection and ritual and autonomously aesthetical tasks got a focal role that increased over time and always was accompanied by emotional events. Behind, the ticking in the deep structure of music of this in cultural-ethical terms totally value-neutral archaic mechanism goes on without pause, contributing to the maintenance of an optimal functional balance in body and mind of the individual, and the group as well.     

Effects of Light Adaptation on the Temporal Resolution of Deep-sea Crustaceans

The effects of light adaptation on flicker fusion frequencywere examined in the photoreceptors of 13 species of deep-seacrustaceans. Light adaptation produced a significant increasein the maximum critical flicker fusion frequency (CFF_max) in7 species—all 6 species of euphausiids in the study, and1 species of oplophorid (Group 1). This is the first exampleof an increase in temporal resolution due to light adaptationin a deep-sea species. In the other six species—2 oplophorids,1 pandalid, 1 pasiphaeid, 1 penaeid and 1 sergestid (Group 2)—lightadaptation had no effect, or resulted in a decrease in the flickerfusion frequency. The mean dark-adapted CFF_max of the Group1 species was significantly higher, and the mean response latencysignificantly lower, than those of the Group 2 species. Possibleexplanations for these differences include the activity andbioluminescence mode of preferred prey items, as well as theretention of larval/juvenile adaptations in adult eyes.     

Genetic explorations of recent human metabolic adaptations: hypotheses and evidence

Since humans and chimpanzees split from a common ancestor over 6 million years ago, human metabolism has changed dramatically. This change includes adaptations to a high-quality diet, the evolution of an energetically expensive brain, dramatic increases in endurance abilities, and capacity for energy storage in white adipose tissue. Human metabolism continues to evolve in modern human populations in response to local environmental and cultural selective forces. Understanding the nature of these selective forces and the physiological responses during human evolution is a compelling challenge for evolutionary biologists. The complex genetic architecture surrounding metabolic phenotypes indicates that selection probably altered allelic frequencies across many loci in populations experiencing adaptive metabolic change to fit their environment. A recent analysis supports this hypothesis, finding that classic selective sweeps at single loci were rare during the past 250 000 years of human evolution. Detection of selective signatures at multiple loci, as well as exploration of physiological adaptation to environment in humans, will require cross-disciplinary collaboration, including the incorporation of biological pathway analysis. This review explores the Thrifty Genotype Hypothesis, high-altitude adaptation, cold-resistance adaptation, and genetic evidence surrounding these proposed metabolic adaptations in an attempt to clarify current challenges and avenues for future progress.     

Indirect Evolution of Hybrid Lethality Due to Linkage with Selected Locus in Mimulus guttatus

Most species are superbly and intricately adapted to the environments in which they live. Adaptive evolution by natural selection is the primary force shaping biological diversity. Differences between closely related species in ecologically selected characters such as habitat preference, reproductive timing, courtship behavior, or pollinator attraction may prevent interbreeding in nature, causing reproductive isolation. But does ecological adaptation cause reproductive incompatibilities such as hybrid sterility or lethality? Although several genes causing hybrid incompatibilities have been identified, there is intense debate over whether the genes that contribute to ecological adaptations also cause hybrid incompatibilities. Thirty years ago, a genetic study of local adaptation to copper mine soils in the wildflower Mimulus guttatus identified a locus that appeared to cause copper tolerance and hybrid lethality in crosses to other populations. But do copper tolerance and hybrid lethality have the same molecular genetic basis? Here we show, using high-resolution genome mapping, that copper tolerance and hybrid lethality are not caused by the same gene but are in fact separately controlled by two tightly linked loci. We further show that selection on the copper tolerance locus indirectly caused the hybrid incompatibility allele to go to high frequency in the copper mine population because of hitchhiking. Our results provide a new twist on Darwin''s original supposition that hybrid incompatibilities evolve as an incidental by-product of ordinary adaptation to the environment.     

Fantastic animals as an experimental model to teach animal adaptation

Background

Science curricula and teachers should emphasize evolution in a manner commensurate with its importance as a unifying concept in science. The concept of adaptation represents a first step to understand the results of natural selection. We settled an experimental project of alternative didactic to improve knowledge of organism adaptation. Students were involved and stimulated in learning processes by creative activities. To set adaptation in a historic frame, fossil records as evidence of past life and evolution were considered.

Results

The experimental project is schematized in nine phases: review of previous knowledge; lesson on fossils; lesson on fantastic animals; planning an imaginary world; creation of an imaginary animal; revision of the imaginary animals; adaptations of real animals; adaptations of fossil animals; and public exposition. A rubric to evaluate the student's performances is reported. The project involved professors and students of the University of Modena and Reggio Emilia and of the "G. Marconi" Secondary School of First Degree (Modena, Italy).

Conclusion

The educational objectives of the project are in line with the National Indications of the Italian Ministry of Public Instruction: knowledge of the characteristics of living beings, the meanings of the term "adaptation", the meaning of fossils, the definition of ecosystem, and the particularity of the different biomes. At the end of the project, students will be able to grasp particular adaptations of real organisms and to deduce information about the environment in which the organism evolved. This project allows students to review previous knowledge and to form their personalities.

     

Adaptivity of social systems: the problem for scientific research

The notion of adaptive evolution of social systems as of a real process of selection of the properties of such systems implies group selection. But strong evidences of effective group selection seem impossible, at least in vertebrates. However, understanding the origin of social systems adaptivity based on individual selection is difficult, as well, without analyzing the proximal mechanisms of the formation of such systems. I suppose that social systems change due to changes of individual features that underlie the proximal mechanisms of the system formation. These features are the characteristics of neurophysiological and hormonal regulatory mechanisms. They are strongly associated with intrinsic biochemical processes and are coded in the genome. Thus, the evolution of social systems is the evolution of their proximal mechanisms. At the same time, the specificity of neurophysiological and hormonal regulation determines not only social interactions, but also the individual behaviour of animals. The most important characteristics of life history, such as the regime of activity, foraging strategy, etc., are strongly affected by the same regulatory mechanisms. This view is useful for understanding the relations combining many features into an integrated and adaptive species-specific life form. I suppose that such forms emerged as evolutionary consequences of changes in regulatory mechanisms adaptive to specific environment. Thus, we have as substantial reasons to discuss adaptations of social systems to ecological features as to discuss ecological features adapted to particular social systems. The species-specificity of regulatory mechanisms is probably based on different kinds of evolutionary choice between the rapidity and the perfection of adaptation, between flexibility and stability, and between sensibility and resistibility. I think that this choice depends largely on the predictability of the environment. The less predictable it is, the more it increases the selective value of sensibility, flexibility, and rapidity of evolution. On the contrary, stable and predictable environment stimulates less rapid but more perfect adaptations. Such choices consolidate in the genome during evolution as specific features of neurophysiological and hormonal regulation systems. These specific features, in their turn, determine ecological, behavioural, and physiological species-specificity. From this point of view, evolutionary changes in social systems can be readily perceived as consequences of the selection of individuals, promoting optimal properties under particular conditional features of regulation systems. The boundary condition for this model is the absence of specificity of the characteristics of regulation systems to different forms of stress. This condition needs to be considered closely.     

Optimierung und ökonomisierung im kontext von evolutionstheorie und phylogenetischer rekonstruktion

The meaning of optimality and economy in phylogenetics and evolutionary biology is discussed. It can be shown that the prevailing concepts of optimality and economy are equivocal as they are not based on strict theoretical positions and as they have a variable meaning in different theoretical contexts. The ideas of optimality and economy can be considered to be identical with the expectation of a relatively simple order in a particular field of study. Although there exists no way of inferring one or several methods of solving scientific problems from the presupposed idea of economy and optimality, a lack of motivation for scientific investigations would result if the concepts of economy and optimality in nature were dropped. By reference to several examples, it is shown that the concepts of optimality and economy are only useful against the background of indispensable theories. If there is a shift from one theory to another, a restriction on the use of these concepts is necessary. Optimality and economy in the sense of operations research in engineering or economical sciences depend on the principle of minimum costs. Both theoretical concepts: technical efficiency in relation to the energy required to run a machine and profit maximation in an economical framework must be shown to be realistic assumptions. In the field of biology processes of optimization and economization are normally discussed under two different views:

1. The concept of economy is used in cases of functional adaptation when the organism makes good use of the building material which is available to fulfill one (or more) functions. The theoretical background must be seen in the energy-consuming aspect of the organism.
2. In evolutionary change and phylogeny ‘economization’ and ‘optimization’ are deduced from the evolutionary theory, and evolution is shown to produce a special kind of biological economy in biological systems (Bock & von Wahlert, 1965). The ‘Okonomie-Prinzip’ or ‘Lesrichtungskriterium’ points out the arguments needed to state a phylogenetic theory and to construct a dendrogram (Peters & Gutmann, 1971).

In every phylogenetic theory concerning the adaptational change in the evolving biological system an explanation for the function of all stages is required. Only those statements should be accepted as phylogenetic theories which are characterized by the demonstration of the process of economization in the functional relations of the evolving organism. The process of adaptation can be determined by the improved chance of some mutants to propagate their genetical information. In this process all functional systems in their interrelations — i.a. mutual dependence — and their relation with the environment add their functional efficiency to the information to be delivered to their progeny, because the more economical biological system in a certain environment will have a better chance to produce offspring. This outcome is affirmed by natural selection which works on all levels of the evolving biological systems (Gutmann & Peters 1973). Nevertheless a judgment about adaptation cannot be taken as a scale of measurement in the phylogenetic process. The conditions in the organism itself and in the environment or in the organic system alone can change in so profound a manner that the marginal conditions of the earlier stages of the process of adaptation are not the same as in the derived ones. During phylogenetic change of the evolving organism the selective strains are also continuously changing. As a consequence no state or invariant concept of economy can cover the different stages of the phylogenetic process. The pragmatical meaning of the theoretical consideration is substantiated by the example of the hydrostatic skeleton theory in which the chordates are derived from metameric worms with a fluid skeleton. Herrn Professor Dr. P. Dullemeijer sind die Verfasser für kritische Lektüre und wertovolle Hinweise zu Dank verpflichtet.     

Microbial selection strategies that enhance the likelihood of developing commercial biological control products

Research interest in utilizing microorganisms to create a microbial environment suppressive to plant pathogens has increased exponentially in recent years. Despite intense interest in developing biological control agents, relatively few antagonists have achieved ‘commercial product’ status. The fact that such a small proportion of active laboratory antagonists are developed into biological control products is partly due to several features common to microbial selection strategies that are widely utilized to obtain putative biological control agents: (a) relatively few candidate microorganisms are tested; (b) microbes are selected based on the results of an assay that does not replicate field conditions; and (c) the amenability of microbes to commercial development is excluded as a selection criterion. Selection strategies that enhance the likelihood of developing commercial biological control products are described. These include making appropriate choices regarding the pathosystem for biological control, the method of microbe isolation, and the method of isolate characterization and performance evaluation. A model system of developing a biological control product active against Gibberella pulicaris (Fries) Sacc. (anamorph: Fusarium sambucinum Fuckel), the primary causal agent of Fusarium dry rot of stored potatoes, is used to illustrate the proposed selection strategy concepts. The crucial importance and methodology is described, of selecting strains with enhanced potential for commercial development based on a strain exhibiting both favorable growth kinetics and bioefficacy when grown in commercially feasible liquid media. Received 06 February 1997/ Accepted in revised form 29 May 1997