Evolution of cyclic sexual reproduction under host-parasite interactions

Alternate changes of sexual and asexual generations was studied theoretically by numerical analyses, from the viewpoint of host-parasite infective interactions. The host species was considered a diploid organism, characterized by two loci determining parasite resistance type and sexual strategy, between which a linkage exists to a certain degree. The sexual reproductive strategy was assumed to be determined by three alleles: asexual, complete sexual and cyclic sexual alleles. The effect of the parasites was represented by decreasing fitness functions of each host-type frequency. Numerical analyses of various linkages between genes and frequency dependence of host fitness revealed that the dynamics varied depending on whether the cyclic sexual allele was dominant or recessive against the complete sexual allele. When the cyclic sexual allele was dominant, the cyclic sexual strain persisted under intermediate frequency dependence of host fitness. On the other hand, when the cyclic sexual allele was recessive, it always tended to spread and to be maintained in the population. In such situations, both complete and cyclic sexual strains coexist, but the asexual strain is lost.     

Reassessment of the role of phospholipids in sexual reproduction by sterol-auxotrophic fungi.

Several genera of oomycete fungi which are incapable of de novo sterol synthesis do not require these compounds for vegetative growth. The requirement for an exogenous source of sterols for sexual reproduction by several members of the Pythiaceae has been questioned by reports of apparent induction and maturation of oospores on defined media supplemented with phospholipids in the absence of sterols. A more detailed examination of this phenomenon suggested that trace levels of sterols in the inoculum of some pythiaceous fungi act synergistically with phospholipid medium supplements containing unsaturated fatty acid moieties to induce oosporogenesis. Phospholipid analysis of one species, Pythium ultimum, suggested that only the fatty acid portion of the exogenous phospholipid is taken up by the fungus. Enrichment of the phospholipid fraction of total cell lipid of P. ultimum with unsaturated fatty acids promoted oospore induction, and enhanced levels of unsaturated fatty acids in the neutral lipid fraction increased oospore viability. For some pythiaceous fungi, the levels of sterols required for the maturation of oospores with appropriate phospholipid medium supplementation suggest that these compounds are necessary only for the sparking and critical domain roles previously described in other fungi.     

Spermatogenesis in Hydra oligactis. II. How temperature controls the reciprocity of sexual and asexual reproduction

Hydra oligactis undergo two mutually exclusive modes of reproduction: at warm temperatures (18-22 degrees C) animals reproduce asexually by budding, while at cold temperatures (10-12 degrees C) gamete differentiation occurs. Using a monoclonal antibody which is specific for cells of the sperm lineage, it was discovered that under conditions where sperm differentiation does not occur (18-22 degrees C), cells continually enter the sperm pathway but progression down the pathway is prematurely halted, effectively blocking the production of sperm. To elucidate the mechanism by which completion of sperm differentiation is controlled, the cell cycle times of interstitial cells entering the sperm pathway at both the restrictive (18 degrees C) and permissive (10 degrees C) temperatures were examined. It was envisaged that at the restrictive temperature the cell cycle times of committed cells would lengthen as they proceeded down the pathway, leading to dilution and eventual loss of cells at later stages of sperm differentiation. This did not occur. Although cells of the sperm lineage were found overall to divide more slowly at 18 degrees C than at 10 degrees C, at both temperatures the cell cycle times shortened as cells proceeded further down the pathway, making a dilution mechanism untenable. The effect of high temperature on the survival of cells was then tested by subjecting animals to a heat shock. Within 12 hr of the increase in temperature, the total number of sperm lineage interstitial cells dropped 10-fold while the total numbers of epithelial and somatic interstitial cells remained virtually unchanged. A distinct consequence of this cell loss was the disappearance of cells furthest down the sperm pathway. It is proposed that as cells move down the sperm pathway, they become increasingly sensitive to high temperature which adversely affects their survival; the higher the temperature, the earlier in the pathway cells die. The lethal effect is abolished by lowering the temperature, allowing sperm differentiation to continue to completion. The possible adaptive advantages of temperature controlling gametogenesis are discussed.     

The role of topoisomerase II in drug resistance.

The conventional laboratory approach to study the mechanisms of drug resistance has been the selection of drug-resistant cell lines by continuous exposure to cytotoxic agents. Such lines, which are selected for resistance to a single agent, frequently display cross-resistance to a number of cytotoxic agents that are unrelated in both structure and proposed mechanism of action. Multidrug-resistant cells display reduced drug accumulation, which is the result of overexpression of a surface glycoprotein (P170). Although resistance to multiple antitumor agents is a common clinical problem in the treatment of cancer, the precise role of the P-glycoprotein-mediated mechanism in human tumors remains to be established. Many alterations in multidrug-resistant cells selected in vitro have been identified. The concomitant expression of multiple phenotypic differences, which appear to be favored by continued and prolonged drug exposure, makes analysis of critical individual resistance pathways more difficult. However, multiple factors may also be involved in the development of clinical resistance. Recent studies have identified alterations in DNA topoisomerase II activity and function as an alternative mechanism that contributes to the multidrug-resistance phenomenon or is responsible for a different type of drug resistance. The precise nature of these changes remains unclear. Available evidence supports the view that expression of the enzyme is an important determinant of cell sensitivity to DNA topoisomerase poisons, but that other changes involved in regulation of enzyme function and/or in the cellular processing of drug-induced DNA damage may be critical in determining the differential pattern of cell response to antitumor agents.     

The evolution of sexual reproduction as a repair mechanism part II. Mathematical treatment of the wheel model and its significance for real systems

The dynamics of populations of self-replicating, hierarchically structured individuals, exposed to accidents which destroy their sub-units, is analyzed mathematically, specifically with regard to the roles of redundancy and sexual repair. The following points emerge from this analysis: 1. A population of individuals with redundant sub-structure has no intrinsic steady-state point; it tends to either zero or infinity depending on a critical accident rate alpha c. 2. Increased redundancy renders populations less accident prone initially, but population decline is steeper if alpha is greater than a fixed value alpha d. 3. Periodic, sexual repair at system-specific intervals prevents continuous decline and stabilizes the population insofar as it will now oscillate between two fixed population levels. 4. The stabilizing sexual interval increases with increased complexity provided this is accompanied by appropriate levels of redundancy. 5. The model closely simulates the dynamics of heterosis effects. 6. Repair fitness is a population fitness: the chance of an individual being repaired is a function of the statistical make-up of the population as a whole at the particular period. Populations living at alpha greater than alpha c either engage in sexual repair at the appropriate time or they die out. 7. The mathematical properties of the model illustrate mechanisms which possibly played a role in the evolution of a mortal soma in relation to sexual reproduction.     

Horizontal gene transfer as a possible evolutionary predecessor of sexual reproduction

Computer modeling shows that high rates of lateral gene transfer (LGT), combined with homologous recombination, enhance selection efficiency, thus making it possible for organisms to acquire larger genomes without irreversible growth of mutation load. In prokaryotes, the optimal (high) rate of LGT cannot evolve because of the “suicidal effect” of modifiers, i.e., alleles that enhance LGT are systematically replaced by alleles that hinder LGT and never vice versa. Therefore, the latter alleles spread despite the fact that they reduce fitness of organisms and populations. This “conflict of interests” can be resolved via evolution of whole-genome reciprocal recombination, which effectively removes the basis for propagation of “selfish” modifiers. Eukaryotic sexual reproduction (amphimixis) probably appeared as the final result of evolution of mechanisms of LGT driven by selection for high interindividual recombination rate. Such selection could have been facilitated at the early stages of eukaryogenesis due to an increase in genome size and high mutation rate caused by higher O₂ concentration and invasion of group II introns.     

The evolution of sexual reproduction as a repair mechanism. Part I. A model for self-repair and its biological implications

Summary The theory is presented that the sexual process is a repair mechanism which maintains redundancy within the sub-structure of hierarchical, self-reproducing organisms. In order to keep the problems within mathematically tractable limits (see Part II), a simple model is introduced: a wheel with 6 spokes, 3 of them vital and 3 redundant, symbolizes the individual (cell or organism). Random accidents destroy spokes; the wheels replicate at regular cycles and engage periodically in pairing and repair phases during which missing spokes are copy-reproduced along the intact spokes of the partner wheel.The hierarchical structure of such a system is analysed and an autonomous unit is defined: this is the unit of minimal hierarchical complexity which is capable of perpetuating autonomously all higher and all lower levels of the hierarchy; this is the central unit of selection.Four basic, physical parameters are isolated which determine the essential features of any eucaryotic life cycle: 1. The number of levels of the hierarchy (unicellular, multicellular, colonial, etc.); 2. the relation between the phases of replication (asexual generations) and repair (sexual generations); 3. the duration of potential repair (haplo- diplo-phases); 4. the position of the sexual partners within the hierarchy (selfing, monecy, dioecy, reproductive individuals within colonies, etc.).The evaluation of fitness components is considered in relation to trends of reproductive patterns in evolution.     

In vitro fertilization as a tool for investigating sexual reproduction of angiosperms

In vitro fertilization (IVF) of isolated male and female gametes of flowering plants was first accomplished in the last decade. Successful isolation of male and female gametes, and culturing of in vitro zygotes to form new plants, is a prelude to the use of IVF for research into the cellular and molecular control of fertilization in higher plants and its application as a tool in biotechnology. Genes unique to male and female gametes and zygotes of higher plants, although currently incompletely characterized, are expected to permit direct molecular dissection of fertilization. By applying IVF and microculture to zygotes and endosperm obtained by both in vivo and in vitro methods, newly activated fusion products may be observed and manipulated in media where they are directly accessible to the techniques of molecular cell biology. IVF and zygote culture may also offer potential for creating new hybrid plants by fusing isolated gametes from different species to produce unique zygotes and ultimately plants that would be impossible to obtain using typical crossing techniques. Transformation and regeneration frequencies using IVF may also be high enough to avoid the necessity of adding controversial antibiotic and herbicide resistant genes to screen transformed products. This review describes advances using IVF in plant sexual reproduction and discusses its potential in the genetic improvement of flowering plants.     

A mathematical model for the phase of sexual reproduction in monogonont rotifers

Recently, the optimal sex allocation in monogonont rotifers is studied in [1], and, as a closely related question, the relative frequencies of the relevant types of mictic females. The authors focus on the evolution of the age at which young mictic females lose their fertilization susceptibility and they address the threshold age of fertilization that maximizes resting egg production. Assuming that a stationary population is achieved, with stable age distribution, they obtain their results, without knowing the stationary population. Our aim is to study this problem in the framework of the theory of nonlinear age-dependent population dynamics developed by G. F. Webb in [13], which is more appropriate from the mathematical point of view and permits to us to obtain analytically the stationary population and consequently it is analytically shown that a threshold age of fertilization equal to the age of maturation is not an ESS, despite the fact that then the production of resting eggs is maximum, which has been obtained by simulation in [1]. Received: 30 June 1998 / Revised version: 17 May 1999 / Published on line: 5 May 2000     

Evolution of sexual mimicry in the orchid subtribe orchidinae: the role of preadaptations in the attraction of male bees as pollinators

Background  

Within the astonishing diversity of orchid pollination systems, sexual deception is one of the most stunning. An example is the genus Ophrys, where plants attract male bees as pollinators by mimicking female mating signals. Unsaturated hydrocarbons (alkenes) are often the key signal for this chemical mimicry. Here we investigate the evolution of these key compounds within Orchidinae by mapping their production in flowers of selected species onto their estimated phylogeny.     

On the unfitness of natural selection to explain sexual reproduction, and the difficulties that remain.

In its essence, the explanatory potential of the theory of natural selection is based on the iterative process of random production and variation, and subsequent non-random, directive selection. It is shown that within this explanatory framework, there is no place for the explanation of sexual reproduction. Thus in Darwinistic literature, sexual reproduction - one of nature's most salient characteristics - is often either assumed or ignored, but not explained. This fundamental and challenging gap within a complete naturalistic understanding of living beings calls for the need of a cybernetic account for sexual reproduction, meaning an understanding of the dynamic and creative potential of living beings to continuously and autonomously produce new organisms with unique and specific constellations.     

Concanavalin A binding to Chlamydomonas eugametos flagellar proteins and its effect on sexual reproduction

The relative amounts of Concanavalin A (Con A) bound by gamete and vegetative flagella of both mating types (mt ⁺ and mt ^-) of Chlamydomonas eugametos were determined using ¹²⁵I-Con A. Con A agglutinated all cell types by cross-linking their flagella in a random manner. No correlation was found between the extent of Con A-binding and Con A-mediated isoagglutination. Con A inhibited the sexual interaction between gametes at various levels. In mt ⁺ gametes it blocked sexual agglutination, whereas in mt ^- gametes it prevented papillar fusion. By SDS-gel electrophoresis nine Con A-binding components were found to be present in flagella. However, it was not possible to allocate a role in sexual agglutination to any of these components since they were present in all cell types, including vegetative cells which are not able to sexually agglutinate.Abbreviations Con A concanavalin A - SDS sodium dodecyl sulphate - TB Tris buffer - PBS phosphate buffered saline - HRP horse radish peroxidase - SEM scanning electron microscope - PAS periodic acid Schiff     

A peculiar method of sexual reproduction in certain new members of the Chlamydomonadaceae

Summary In the course of a study of Algae from Indian soils two new species of Chlamydomonas (C. Iyengari, C. indica) and a new species of Carteria (C. eugametos) were observed which are distinctive in the fact that their gametes conjugate by their posterior ends. Diagnoses of the new species are given. The gametes are provided with membranes. One of the fusing gametes receives the contents of the other, and the membrane of the active gamete, which has fused with that of the recipient gamete, forms a loose envelope around the zygote. The zygotes retain only the flagella of the recipient gamete. They are larger than the vegetative cells and may remain motile for some days. They frequently divide without a resting period, although zygospores were formed in old cultures of two of the species. Germination of these zygospores was observed.The author is indebted to Prof. F. E. Fritsch, F. R. S. for advice and guidance in the course of this investigation.     

A physical origin for functional domain structure in nucleic acids as evidenced by cross-linking entropy: II.

In Part I, cross-linking entropy (CLE) was proposed as a mechanism that limits the size of functional domains of RNA. To test this hypothesis, the theory is developed into an RNA secondary structure prediction filter which is applied to nearest-neighbor secondary structure (NNSS) algorithms that utilize a free energy (FE) minimization strategy. (The NNSS strategies are also referred to as the dynamic programming algorithm in the literature.) The cross-linking entropy for RNA is derived from a generalized Gaussian polymer chain model where the entropic contributions caused by the formation of base pairs (stacking) in RNA are analysed globally. Local entropic contributions are associated with the freezing out of degrees of freedom in the links. Both global and local entropic effects are strongly influenced by the persistence length. The cross-linking entropy provides a physical origin for the size of functional domains in long nucleic acid sequences and may go further to explain as to why the majority of the domain regions in typical sequences tend to be less than 600 nucleotides in length. In addition, improvements were observed in the "best guess" predictive capacity over NNSS prediction strategies. The thermodynamic distribution is more representative of the expected structures and is strongly governed by such physical parameters as the persistence length and the excluded volume. The CLE appears to generalize the tabulated penalties used in NNSS algorithms. The principal parameter influencing this entropy is the persistence length. The model is shown to accomodate a variable persistence length and is capable of describing the folding dynamics of RNA. A two-state kinetic model based on the CLE principle is used to help elucidate the folding kinetics of functional domains in the group I introns.     

Character compatibility of molecular markers to distinguish asexual and sexual reproduction

Particularly in polyploids, the potential of the high variability of dominant markers such as random amplified polymorphic DNA fragments (RAPDs) and amplified fragment length polymorphisms (AFLPs) in population genetic studies and analysis of breeding systems is reduced due to their dominant nature. In contrast, the criterion of character compatibility is hindered neither by dominance nor by polyploidy as allelic interpretation is not necessary. Character compatibility, which can be used to detect events of genetic exchange (or recombination), is particularly informative if these events are expected to be rare such as in taxa with extensive vegetative reproduction or apomixis. Binary unordered characters such as presence and absence of anonymous DNA markers are incompatible if all four pairwise combinations of character states are present among the individuals studied. Because incompatible character state distributions defy any progenitor–derivative relationship among individuals, they provide strong evidence for genetic exchange. Both the absolute number of incompatible character combinations and the probability of compatibility can be used as a measure of incompatibility. Although these measures may not directly relate to the frequency of genetic exchange, they provide a useful tool to heuristically explore data sets. The most commonly used input for multivariate analyses and analysis of molecular variance in population genetic studies of (dis)similarity of marker distributions are amalgamates of mutation and recombination. Character compatibility can be used to complement these traditional methods of analysis. Advantages and disadvantages of character incompatibility relative to multilocus analysis of modes of reproduction and population genetics are demonstrated with data from RAPDs, isozymes, and restriction fragment length polymorphisms (RFLPs) of the nuclear ribosomal and chloroplast genome.