Evolution as resistance to entropy. I. Mechanisms of species homeostasis

The idea is discussed that the common output of any evolution is creation of the entities that are increasingly resistant to further evolution. The moving force of evolution is entropy, the tendency to disorder. This general aspiration for chaos is a cause of the mortality of organisms and species, however, being prerequisite for any movement, it creates (by chance) novelties, which may occur (by chance) more resistant to further decay and thus survive. The surviving of those who survive is the most general principle of evolution discovered by Darwin for particular case of biological evolution. The second law of thermodynamics states that our Universe is perishing but its ontology is such that it creates resistance to destruction. The evolution is a history of this resistance. Not only those who die do not survive but also those who evolve. The entities that change (evolve) rapidly disappear rapidly and by this reason they are not observed among both the fossils and now-living organisms. We know only about long-living species. All the existing organisms are endowed with an ability to resist other changing. The following main achievements of the species homeostasis are discussed: high fidelity of DNA replication and effective mechanisms of DNA repair; diploidy; normalizing selection; truncated selection; heterozygote superiority; ability to change phenotype adaptively without changing genotype; parental care and the K-strategy of reproduction; behavior that provides independence of the environment. The global resistance of the living systems to entropy is provided the state that all the essential in biology is determined not by physical-chemical interactions but could semantic rules. A conception of "potential zygotic information" that determines the rules of ontogenesis is proposed. A zygote does not contain this information in explicit form. It is created de novo step by step during ontogenesis and it could not be decoded beforehand. The experimental data on the adaptive mutagenesis and the relevant hypothesis are discussed. It is concluded that the special mechanisms for speeding-up of evolution as created by evolution are impossible conceptually.     

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.     

Zygospores as evidence of sexual reproduction in the genus Orphella

The presence of zygospores in the genus Orphella is newly described. We found zygospores in three species of the genus, O. catalaunica, O. coronata and O. helicospora, which are all the species of the genus known from the Iberian territory. Zygospores are associated with a heterothallic conjugating sexual process in O. coronata, whereas in O. catalaunica and O. helicospora, they form homothallically. In all instances, zygospores are consistently associated with an organized pattern of sterile cells, forming structures comparable to those present with asexual trichospores. We compare the ontogeny of Orphella zygospores with that found in the harpellid Genistellospora homothallica and discuss the possible close relationship of Orphella with Kickxellales (Zygomycetes). We report O. coronata in Spain for the first time, replacing all previous records of O. haysii. Results are supported with line drawings and photographs.     

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.     

毛红椿天然种群有性繁殖适合度及其繁殖更新

以九连山国家自然保护区毛红椿4个天然种群为研究对象,于2006—2016年调查毛红椿有性繁殖和自然更新的情况,分析其繁殖适合度系数和个体水平适合度.结果表明:九连山毛红椿各地理种群结实单株的初始数量较少(3~9株),经70余年的繁衍发育,现存结实单株的数量仅2~10株,且来源于原始单株或子1~2代.不同种群间有性繁殖差异显著,但结实能力呈逐年下降的趋势;随群落发育成熟,土壤种子库保存与种子萌发的失利,导致能正常生长发育成熟的林木数量近乎为0.毛红椿最佳性成熟年龄为40年,早期优势明显,适合度系数早期快速增长,为2.0~2.8,但急剧减少至0.3~0.5,之后较平稳减速至近乎于0;种群间个体有性繁殖适合度(0~14株·cm2)存在较显著差异,但均较低,甚至低至0;以现有繁殖率计算,有性繁殖与更新的适合度实测值均远低于预估值.总之,受有性繁殖遗传适合度低的影响,不同种群间有性繁殖与更新能力趋同衰退;个体有性繁殖适合度进一步降低且面临更高的投资风险,现有繁殖体系因此失衡并趋于恶化.建议开展繁殖交配、授粉结实及遗传多样性评价研究,同时人为干预林分环境,于结实期清理林地枯落物,在幼苗至幼树生长过渡期适当疏伐.     

Glucuronidation of type II arabinogalactan polysaccharides function in sexual reproduction of Arabidopsis

Arabinogalactan proteins (AGPs) are complex, hyperglycosylated plant cell wall proteins with little known about the biological roles of their glycan moieties in sexual reproduction. Here, we report that GLCAT14A, GLCAT14B, and GLCAT14C, three enzymes responsible for the addition of glucuronic acid residues to AGPs, function in pollen development, polytubey block, and normal embryo development in Arabidopsis. Using biochemical and immunolabeling techniques, we demonstrated that the loss of function of the GLCAT14A, GLCAT14B, and GLCAT14C genes resulted in disorganization of the reticulate structure of the exine wall, abnormal development of the intine layer, and collapse of pollen grains in glcat14a/b and glcat14a/b/c mutants. Synchronous development between locules within the same anther was also lost in some glcat14a/b/c stamens. In addition, we observed excessive attraction of pollen tubes targeting glcat14a/b/c ovules, indicating that the polytubey block mechanism was compromised. Monosaccharide composition analysis revealed significant reductions in all sugars in glcat14a/b and glcat14a/b/c mutants except for arabinose and galactose, while immunolabeling showed decreased amounts of AGP sugar epitopes recognized by glcat14a/b and glcat14a/b/c mutants compared with the wild type. This work demonstrates the important roles that AG glucuronidation plays in Arabidopsis sexual reproduction and reproductive development.     

Evolution: sociality as a driver of unorthodox reproduction

An unusual reproductive system was discovered in desert ants, in which daughter queens are produced asexually via parthenogenesis, whereas workers develop from hybrid crosses between genetically divergent lineages. The system appears to be doomed to extinction.     

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.     

A centrosomal theory of the short term evolutionary maintenance of sexual reproduction

A new mode of inheritance is postulated in which a sexual offspring receives a contribution from each parent and selects the better to pass on to its own offspring. This could provide a simple advantage to sex over a sex whose magnitude is shown to be of the order of a doubling of fitness in each generation, large enough to cancel the twofold cost of sex. This possible advantage to sex can be realized only if a cell component is in fact inherited in this selectively ambiguous way. No such component is known of, but the eukaryotic centrosome is a possible candidate. The possibility is discussed that the centrosome contains an obligatorily non-digital replicator which has an essential function in the initiation of microtubules. If this theory is true, it has the capacity to apply as widely as sex is found, and it would rescue theories of the long-term maintenance of sex from the necessity to provide a twofold advantage in each generation. If false, the theory will soon be disproved.     

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     

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.     

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.