How many processes are responsible for phenotypic evolution?

SUMMARY In addressing phenotypic evolution, this article reconsiders natural selection, random drift, developmental constraints, and internal selection in the new extended context of evolutionary developmental biology. The change of perspective from the "evolution of phenotypes" toward an "evolution of ontogenies" (evo-devo perspective) affects the reciprocal relationships among these different processes. Random drift and natural selection are sibling processes: two forms of post-productional sorting among alternative developmental trajectories, the former random, the latter nonrandom. Developmental constraint is a compound concept; it contains even some forms of natural ("external" and "internal") selection. A narrower definition ("reproductive constraints") is proposed. Internal selection is not a selection caused by an internal agent. It is a form of environment-independent selection depending on the level of the organism's internal developmental or functional coordination. Selection and constraints are the main deterministic processes in phenotypic evolution but they are not opposing forces. Indeed, they are continuously interacting processes of evolutionary change, but with different roles that should not be confused.     

The enigma of frequency-dependent selection

Frequency-dependent selection is so fundamental to modern evolutionary thinking that everyone `knows' the concept. Yet the term is used to refer to different types of selection. The concept is well defined in the original context of population genetics theory, which focuses on short-term evolutionary change. The original concept becomes ambiguous, however, when used in the context of long-term evolution, where density dependence becomes essential. Weak and strong frequency dependence, as distinguished in this article, refer to two very different forms of selection.     

Crowdedness Mediates the Effect of Social Identification on Positive Emotion in a Crowd: A Survey of Two Crowd Events

Exposure to crowding is said to be aversive, yet people also seek out and enjoy crowded situations. We surveyed participants at two crowd events to test the prediction of self-categorization theory that variable emotional responses to crowding are a function of social identification with the crowd. In data collected from participants who attended a crowded outdoor music event (n = 48), identification with the crowd predicted feeling less crowded; and there was an indirect effect of identification with the crowd on positive emotion through feeling less crowded. Identification with the crowd also moderated the relation between feeling less crowded and positive emotion. In data collected at a demonstration march (n = 112), identification with the crowd predicted central (most dense) location in the crowd; and there was an indirect effect of identification with the crowd on positive emotion through central location in the crowd. Positive emotion in the crowd also increased over the duration of the crowd event. These findings are in line with the predictions of self-categorization theory. They are inconsistent with approaches that suggest that crowding is inherently aversive; and they cannot easily be explained through the concept of ‘personal space’.     

The dynamical attainability of ESS in evolutionary games

In this paper, the attainability of ESS of the evolutionary game among n players under the frequency-independent selection is studied by means of a mathematical model describing the dynamical development and a concept of stability (strongly determined stability). It is assumed that natural selection and small mutations cause the phenotype to change gradually in the direction of fitness increasing. It is shown that (1) the ESS solution is not always evolutionarily attainable in the evolutionary dynamics, (2) in the game where the interaction between two species is completely competitive, the Nash solution is always attainable, and (3) one of two species may attain the state of minimum fitness as a result of evolution. The attainability of ESS is also examined in two game models on the sex ratio of wasps and aphids in light of our criterion of the attainability of ESS.     

Epizootic of toxoplasmosis among the crowned pigeon, Goura cristata Pallas and Goura victoria Frazer, of the Antwerp Zoo

The authors report an epizootic form of toxoplasmosis observed among the crowned pigeons (Goura cristata Pallas and Goura victoria Frazer). The evolution of toxoplasma from pigeons is described and the epizootiological analysis of two epizootic forms of toxoplasmosis encountered in Antwerp Zoo are compared.     

松科早期球果研究的历史和现状

本文主要对早白垩世到老第三纪渐新世这段时间的松科化石球果的研究作了系统回顾和总结。松科化石球果的研究经历了一个由分散、幼稚到系统、成熟的发展过程,现在将松科化石球果分为三个属：Pityostrobus、Pseudoaraucaria 及Pinus。其中Pseudoaraucaria及Pinus为两个自然属,Pseudoaraucaria已经绝灭。Pityostrobus作为一个器官属,包括那些在松科中分类位置不明的球果。对松科化石球果进行细致的外部形态及内部结构的研究主要目的在于演化关系的探讨上：对于Pinus,主要探讨这个属内各个亚属、组、亚组之间的演化关系;对于Pityostrobus和Pseudoaraucaria,通过绝灭类型的分析,探索松科内各个类群(包括绝灭的)的演化关系进而完成进化树是研究的主要方向,现在这方面已有了Pinus中心说及Pseudoaraucaria分支说两个假说。     

松科早期球果研究的历史与现状

本文主要对早白垩世到老第三纪渐新世这段时间的松科化石球果的研究作了系统回顾和总结。松科化石球果的研究经历了一个由分散、幼稚到系统、成熟的发展过程,现在将松科化石球果分为三个属：Ｐｉｔｙｏｓｔｒｏｂｕｓ、Ｐｓｅｕｄｏａｒａｕｃａｒｉａ及Ｐｉｎｕｓ。其中Ｐｓｅｕｄｏａｒａｕｃａｒｉａ及Ｐｉｎｕｓ为两个自然属,Ｐｓｅｕｄｏａｒａｕｃａｒｉａ已经绝灭。Ｐｉｔｙｏｓｔｒｏｂｕｓ作为一个器官属,包括那些在松科中分类位置不明的球果。对松科化石球果进行细致的外部形态及内部结构的研究主要目的在于演化关系的探讨上：对于Ｐｉｎｕｓ,主要探讨这个属内各个亚属、组、亚组之间的演化关系;对于Ｐｉｔｙｏｓｔｒｏｂｕｓ和Ｐｓｅｕｄｏａｒａｕｃａｒｉａ,通过绝灭类型的分析,探索松科内各个类群（包括绝灭的）的演化关系进而完成进化树是研究的主要方向,现在这方面已有了Ｐｉｎｕｓ中心说及Ｐｓｅｕｄｏａｒａｕｃａｒｉａ分支说两个假说。     

The sympatric breeding of Common and Scottish Crossbills Loxia curvirostra and L. scotica and the evolution of crossbills

Within the strongly polytypic Red Crossbill L. curvirostra there are many reports of two or more 'subspecies' nesting sympatrically, without interbreeding. This 13-year study examines one such case, in Scotland, where an endemic form is resident and another was thought to occur after irruptions from its main range in continental Europe. Both forms were present in the study area every year; sympatric breeding was proved in 9 years and probably occurred in the other four. There was no suggestion of interbreeding and the Scottish form should be treated as a separate species, L. scotica , the Scottish Crossbill. This is the only endemic species of bird in Britain and one of very few European endemics. The evolution of crossbills in Europe is discussed in the light of recent palynological evidence and the taxonomic status of the Parrot Crossbill L. pytyopsittacus is re-examined.     

The involvement of two cdc2-related kinases (CRKs) in Trypanosoma brucei cell cycle regulation and the distinctive stage-specific phenotypes caused by CRK3 depletion

Cyclin-dependent protein kinases are among the key regulators of eukaryotic cell cycle progression. Potential functions of the five cdc2-related kinases (CRK) in Trypanosoma brucei were analyzed using the RNA interference (RNA(i)) technique. In both the procyclic and bloodstream forms of T. brucei, CRK1 is apparently involved in controlling the G(1)/S transition, whereas CRK3 plays an important role in catalyzing cells across the G(2)/M junction. A knockdown of CRK1 caused accumulation of cells in the G(1) phase without apparent phenotypic change, whereas depletion of CRK3 enriched cells of both forms in the G(2)/M phase. However, two distinctive phenotypes were observed between the CRK3-deficient procyclic and bloodstream forms. The procyclic form has a majority of the cells containing a single enlarged nucleus plus one kinetoplast. There is also an enhanced population of anucleated cells, each containing a single kinetoplast known as the zoids (0N1K). The CRK3-depleted bloodstream form has an increased number of one nucleus-two kinetoplast cells (1N2K) and a small population containing aggregated multiple nuclei and multiple kinetoplasts. Apparently, these two forms have different mechanisms in cell cycle regulation. Although the procyclic form can be driven into cytokinesis and cell division by kinetoplast segregation without a completed mitosis, the bloodstream form cannot enter cytokinesis under the same condition. Instead, it keeps going through another G(1) phase and enters a new S phase resulting in an aggregate of multiple nuclei and multiple kinetoplasts in an undivided cell. The different leakiness in cell cycle regulation between two stage-specific forms of an organism provides an interesting and useful model for further understanding the evolution of cell cycle control among the eukaryotes.     

Epigenetic mechanisms of character origination

The close mapping between genotype and morphological phenotype in many contemporary metazoans has led to the general notion that the evolution of organismal form is a direct consequence of evolving genetic programs. In contrast to this view, we propose that the present relationship between genes and form is a highly derived condition, a product of evolution rather than its precondition. Prior to the biochemical canalization of developmental pathways, and the stabilization of phenotypes, interaction of multicellular organisms with their physicochemical environments dictated a many-to-many mapping between genomes and forms. These forms would have been generated by epigenetic mechanisms: initially physical processes characteristic of condensed, chemically active materials, and later conditional, inductive interactions among the organism's constituent tissues. This concept, that epigenetic mechanisms are the generative agents of morphological character origination, helps to explain findings that are difficult to reconcile with the standard neo-Darwinian model, e.g., the burst of body plans in the early Cambrian, the origins of morphological innovation, homology, and rapid change of form. Our concept entails a new interpretation of the relationship between genes and biological form.     

On the dimensionality of ecological stability

Ecological stability is touted as a complex and multifaceted concept, including components such as variability, resistance, resilience, persistence and robustness. Even though a complete appreciation of the effects of perturbations on ecosystems requires the simultaneous measurement of these multiple components of stability, most ecological research has focused on one or a few of those components analysed in isolation. Here, we present a new view of ecological stability that recognises explicitly the non‐independence of components of stability. This provides an approach for simplifying the concept of stability. We illustrate the concept and approach using results from a field experiment, and show that the effective dimensionality of ecological stability is considerably lower than if the various components of stability were unrelated. However, strong perturbations can modify, and even decouple, relationships among individual components of stability. Thus, perturbations not only increase the dimensionality of stability but they can also alter the relationships among components of stability in different ways. Studies that focus on single forms of stability in isolation therefore risk underestimating significantly the potential of perturbations to destabilise ecosystems. In contrast, application of the multidimensional stability framework that we propose gives a far richer understanding of how communities respond to perturbations.     

Biology Needs Information Theory

Communication is an important feature of the living world that mainstream biology fails to adequately deal with. Applying two main disciplines can be contemplated to fill in this gap: semiotics and information theory. Semiotics is a philosophical discipline mainly concerned with meaning; applying it to life already originated in biosemiotics. Information theory is a mathematical discipline coming from engineering which has literal communication as purpose. Biosemiotics and information theory are thus concerned with distinct and complementary possible meanings of the word ‘communication’. Since literal communication needs to be secured so as to enable semantics being communicated, information theory is a necessary prerequisite to biosemiotics. Moreover, heredity is a purely literal communication process of capital importance fully relevant to literal communication, hence to information theory. A short introduction to discrete information theory is proposed, which is centred on the concept of redundancy and its use in order to make sequences resilient to errors. Information theory has been an extremely active and fruitful domain of researches and the motor of the tremendous progress of communication engineering in the last decades. Its possible connections with semantics and linguistics are briefly considered. Its applications to biology are suggested especially as regards error-correcting codes which are mandatory for securing the conservation of genomes. Biology needs information theory so biologists and communication engineers should closely collaborate.     

Phenotypic Novelty in EvoDevo: The Distinction Between Continuous and Discontinuous Variation and Its Importance in Evolutionary Theory

The introduction of novel phenotypic structures is one of the most significant aspects of organismal evolution. Yet the concept of evolutionary novelty is used with drastically different connotations in various fields of research, and debate exists about whether novelties represent features that are distinct from standard forms of phenotypic variation. This article contrasts four separate uses for novelty in genetics, population genetics, morphology, and behavioral science, before establishing how novelties are used in evolutionary developmental biology (EvoDevo). In particular, it is detailed how an EvoDevo-specific research approach to novelty produces insight distinct from other fields, gives the concept explanatory power with predictive capacities, and brings new consequences to evolutionary theory. This includes the outlining of research strategies that draw attention to productive areas of inquiry, such as threshold dynamics in development. It is argued that an EvoDevo-based approach to novelty is inherently mechanistic, treats the phenotype as an agent with generative potential, and prompts a distinction between continuous and discontinuous variation in evolutionary theory.     

Prey-race drives differentiation of biotypes in ant-eating spiders

1.?Disruptive natural selection resulting from specialization on different hosts is recognized as one of the most important driving forces in the diversification of herbivores and parasites. It has been proposed that a similar mechanism could apply to carnivorous predators too, although the evidence is still lacking. 2.?Here, we show that the differentiation of biotypes of specialized ant-eating spiders of the genus Zodarion has probably been induced by prey-shifting. We focused on two forms of one species Z.?styliferum from the Iberian Peninsula that presumably represent ecological races. We conducted geographic, ecological, venom-oriented, reproductive and genetic divergence analysis among multiple populations collected at a number of sites across Portugal and Madeira. 3.?Geographic analysis revealed that the two forms occur in mosaic sympatry. Each form was found to associate in nature with a different ant species in a different habitat. Specifically, the styliferum form hunted predominantly Messor ants, and the extraneum form hunted mainly Camponotus ants. Laboratory experiments revealed that the two forms exhibit a significant preference for attacking focal ants, demonstrating higher paralysis efficiency, and also show different venom composition. Cross-mating of the two forms was significantly less likely than between pairs of the same form, suggesting moderate assortative mating. Phylogenetic analyses indicate low genetic differentiation of the two forms and parallel-repeated evolution of biotypes. 4.?Adaptive prey-shifting correlated with habitat preference are at present the most valid explanations for biotype formation in Zodarion. The speciation of ant-eating Zodarion spiders thus appears to follow a scenario similar to that of host-shifting in parasites and herbivores.     

Design – an inappropriate concept in evolutionary theory*

The concept of accident in evolution refers to causes which are stochastic with respect to selective demands arising from the external environment and acting on the organism, while the concept of design refers to causes which meet the requirement of these selective demands. The condition ‘with respect to selective demands’ is generally forgotten so that evolutionary changes are described as being design modifications. Design is an invalid synonym for adaptation. Further it implies a designer and has been used by some authors since before Darwin to argue that design in organisms demonstrates the existence of a designer and hence a plan. Yet if evolution depends on two simultaneously acting causes, one of which is accidental, then the process of evolution and all attributes of organisms are accidental. The concept of design is inappropriate in biology and should be eliminated from all biological explanations.     

Stability in an annually defaunated estuarine soft-bottom community

Summary The concept of biological stability is so complex that at least six different meanings have been ascribed to it. We propose that one definition, the ability of a system once perturbed to return to its previous state be utilized as a working definition. Using quantitative data collected monthly from a soft-bottom community that undergoes an annual natural catastrophic defaunation coupled with a recently developed analytical technique, we demonstrate the feasability of a working definition and show the existence of stability in the soft-bottom community. The utility of a working definition of stability in the evaluation of disturbance is discussed.A portion of a dissertation (SLS) submitted in partial fulfillment of the Ph.D. degree, Department of Biology, University of South Florida, Tampa, Florida     

Threshold policies control for predator-prey systems using a control Liapunov function approach

The stability of predator-prey models, in the context of exploitation of renewable resources, subject to threshold policies (TP) is studied in this paper using the idea of backstepping and control Liapunov functions (CLF) well known in control theory, as well as the concept of virtual equilibria. TPs are defined and analysed for different types of one and two species predator-prey models. The models studied are the single species Noy-Meir herbivore-vegetation model, in a grazing management context, as well as the Rosenzweig-MacArthur two species predator-prey model, in a fishery management context. TPs are shown to be versatile and useful in managing renewable resources, being simple to design and implement, and also yielding advantages in situations of overexploitation.     

Stable generation of simple forms

It has been suggested by Thom that the mathematical theory of structural stability gives insight into the stable reproduction of form in biological systems. In this paper a process for the generation of some simple forms is described and it is shown that the forms produced are insensitive to perturbations in the production process. The reason for this stability is found in an analysis of the process in terms of the theory of structural stability and in particular in terms of “catastrophes”. Based on the example of our form generating process, we give a general definition of form and define the pattern recognition problem (which is inverse to the generation problem) in the context of that definition.     

Chromosome differentiation of four 2n = 50 chromosomal forms of Turkish mole rat, Nannospalax nehringi

Nannospalax is a genus of blind rodents adapted to living in underground. The species have numerous chromosomal forms in Turkey, and their taxonomic position is still unknown. In this study, 15 mole rats of four different 2n = 50 forms were used; C- and G- banding processes were applied; and a comparison was made accordingly. Karyological results showed that the 2n = 50S form is a new form for Turkish blind mole rats. 2n = 50S form is determined from Andirin (Kahramanmara?) and has NF = 70. The 2n = 50W form, on the other hand, differs from the others with NF = 74 form. C-banding results showed that heterochromatin blocks of all 2n = 50 are different, while only the 2n = 50W form has telomeric heterochromatin blocks. G-banding results, however, displayed homologies and differences among the chromosomal forms. After comparison, we determined that Robertsonian fusion is an efficient force on chromosomal evolution in blind mole rats in Turkey, and that telomeric heterochromatin is a distinctive character for the 2n = 50W form. We suggest that the chromosomal changing mechanism should be independent from climatic peculiarities. These results support the theory that ancestral karyotype should have the largest distribution in a chromosomally variable species.     

Ultrastructure of spore development in <Emphasis Type="Italic">Scutellospora heterogama</Emphasis>

The ultrastructural detail of spore development in Scutellospora heterogama is described. Although the main ontogenetic events are similar to those described from light microscopy, the complexity of wall layering is greater when examined at an ultrastructural level. The basic concept of a rigid spore wall enclosing two inner, flexible walls still holds true, but there are additional zones within these three walls distinguishable using electron microscopy, including an inner layer that is involved in the formation of the germination shield. The spore wall has three layers rather than the two reported previously. An outer, thin ornamented layer and an inner, thicker layer are both derived from the hyphal wall and present at all stages of development. These layers differentiate into the outer spore layer visible at the light microscope level. A third inner layer unique to the spore develops during spore swelling and rapidly expands before contracting back to form the second wall layer visible by light microscopy. The two inner flexible walls also are more complex than light microscopy suggests. The close association with the inner flexible walls with germination shield formation consolidates the preferred use of the term ‘germinal walls’ for these structures. A thin electron-dense layer separates the two germinal walls and is the region in which the germination shield forms. The inner germinal wall develops at least two sub-layers, one of which has an appearance similar to that of the expanding layer of the outer spore wall. An electron-dense layer is formed on the inner surface of the inner germinal wall as the germination shield develops, and this forms the wall surrounding the germination shield as well as the germination tube. At maturity, the outer germinal wall develops a thin, striate layer within its substructure.