The importance of sexual and asexual reproduction in the recent evolution of Allium vineale

In the weedy plant species Allium vineale (wild garlic), individuals may simultaneously produce sexually and asexually derived offspring, by seed and bulbils, respectively. In this study, genetic and genotypic diversity was determined in samples from 14 European A. vineale populations using nuclear (RAPD) and cytoplasmic (PCR-RFLP of cpDNA) markers to investigate the importance of the different reproductive modes. In the whole sample, 77 nuclear multilocus genotypes and four chloroplast haplotypes (chlorotypes) were found. Populations exhibited a high degree of subdivision for nuclear and cytoplasmic markers as estimated from hierarchical F-statistics; at the same time, identical chlorotypes could be found in populations separated by large distances. Genotypic diversity was significantly lower than expected under free recombination in almost all populations, indicating that recruitment into populations is mostly by asexually produced offspring. Nevertheless, within each chlorotype, the distribution of markers from pairs of nuclear loci was incompatible with a purely clonal structure, suggesting that many multilocus genotypes have originated by sexual recombination rather than by mutation within asexual lineages. It is argued that the weedy habit of A. vineale is likely to have favored bulbil reproduction, whereas sexually generated genotypes may have facilitated local adaptation during the species' expansion across Europe.     

Patterns of sexual reproduction in diatoms

Sexual reproduction takes many forms within the diatoms. The variation has been classified by several authors, but in most cases the distinctions between their main categories have depended on the number of gametes produced per gametangium (and thus on how many zygotes per pair of copulating cells), and upon whether fusion is oogamous, anisogamous or isogamous. These classifications are not themselves an adequate basis for taxonomic comparison, which should be based on individual characteristics of the sexual process. Diatoms seem to be primitively oogamous. In araphid pennate diatoms and some raphid diatoms the gametes and gametangia are morphologically alike but physiologically distinct; one gametangium produces active gametes and the other passive ones. This may be the primitive condition in pennate diatoms, providing a link to the oogamy of centrics via the morphological anisogamy of Rhabdonema Kütz.     

Phenotypic variation,sexual reproduction and evolutionary population dynamics

I studied the effects of introducing phenotypic variation into a well-known single species model for a population with discrete, non-overlapping generations. The phenotypes differed in their dynamic behaviour. The analysis was made under the assumption that the population was in an evolutionary stable state. Differences in the timing of the competitive impacts of the phenotypes on each other had a strong simplifying effect on the dynamics. This result could also be applied to competition between species. The effect of sexual reproduction on the dynamics of the population was analysed by assuming the simplest genetic model of one locus with two alleles. Sexual reproduction made the system much more stable in the (mathematical) sense that the number of attractors was reduced and their basins of attraction enlarged. In a dominant system sex tended to increase the frequency of the recessive allele, and in an overdominant system it induced gene frequencies of 1/2. Whether the attractors in the dominant system tended to be simpler or more complex than the attractors in the asexual system depended on the phenotype of the recessive homozygote. The overdominant sexual system tended to have simpler dynamics than the corresponding asexual population. A 2-locus model was used to study whether sexuals can invade an asexual population and vice versa. One locus coded for sexual and asexual reproduction, while the other coded for the dynamics. Enhanced stability through sexual reproduction seemed to be the reason why there was a clear asymmetry favouring sex in this evolutionary context.     

The conservation of redundancy in genetic systems: effects of sexual and asexual reproduction

The relationship between probability of survival and the number of deleterious mutations in the genome is investigated using three different models of highly redundant systems that interact with a threatening environment. Model one is a system that counters a potentially lethal infection; it has multiple identical components that act in sequence and in parallel. Model two has many different overlapping components that provide three-fold coverage of a large number of vital functions. The third model is based on statistical decision theory: an ideal detector, following an optimum decision strategy, makes crucial decisions in an uncertain world. The probability of a fatal error is reduced by a redundant sampling system, but the chance of error rises as the system is impaired by deleterious mutations. In all three cases the survival profile shows a synergistic pattern in that the probability of survival falls slowly and then more rapidly. This is different than the multiplicative or independent survival profile that is often used in mathematical models. It is suggested that a synergistic profile is a property of redundant systems. Model one is then used to study the conservation of redundancy during sexual and asexual reproduction. A unicellular haploid organism reproducing asexually retains redundancy when the mutation rate is very low (0001 per cell division), but tends to lose high levels of redundancy if the mutation rate is increased (001 to 01 per cell division). If a similar unicellular haploid organism has a sexual phase then redundancy is retained for mutation rates between 0001 and 01 per cell division. The sexual organism outgrows the asexual organism when the above mutation rates apply. If they compete for finite resources the asexual organism will be extinguished. Variants of the sexual organism with increased redundancy will outgrow those with lower levels of redundancy and the sexual process facilitates the evolution of more complex forms. There is a limit to the extent that complexity can be increased by increasing the size of the genome and in asexual organisms this leads to progressive accumulation of mutations with loss of redundancy and eventual extinction. If complexity is increased by using genes in new combinations, the asexual form can reach a stable equilibrium, although it is associated with some loss of redundancy. The sexual form, by comparison, can survive, with retention of redundancy, even if the mutation rate is above one per generation. The conservation and evolution of redundancy, which is essential for complexity, depends on the sexual process of reproduction.     

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.     

Arabinogalactan proteins in plant sexual reproduction

Summary Arabinogalactan proteins (AGPs) are a class of plant extracellular-matrix proteins believed to participate in a broad range of processes involving the plant cell surface. They are extremely abundant in female reproductive tissues and in pollen tubes, the haploid male structures that traverse the diploid female reproductive tissues to deliver sperms to the egg cells. The prevalence of AGPs in reproductive tissues has led to speculations that they play significant functional roles ranging from serving as nutrient resources to cell-cell recognition in plant reproduction. Recent research from several laboratories demonstrated functional participation by AGPs in reproductive processes and began to examine the mechanisms underlying these functional roles. An overview of these recent studies will be discussed with a historical perspective as well as with a view towards future studies in establishing the significance of AGPs that, as a class, they have prominent roles in plant sexual reproduction in multiple and diverse ways.     

The Tangled Bank: The maintenance of sexual reproduction through competitive interactions

The maintenance of sexual reproduction is discussed using a model based on the familiar Lotka-Volterra competition equations. Both the equilibrium and the stability conditions that allow a sexual population to resist invasion by a single asexual clone are considered. The equilibrium conditions give results similar to previous models: When the cost of sex, within phenotype niche width, and environmental variance are low, the sexual population coexists with the asexual clone and remains at a high density. However, the asexual clone is never completely excluded. Analysis of the stability conditions shows a different picture: The introduction of an asexual clone considerably reduces the stability of the community. However, owing to its larger total niche width, the sexual population exists partly in a “competitor-free space” where the asexual clone has almost no influence on the outcome of the interactions. Therefore the asexual clone is less stable than the sexual population and has a higher probability of extinction. In contrast, the sexual population does not become extinct, since the extreme phenotypes remain at a stable, though low, density, and the central phenotypes, where stability is low, are recreated every generation through recombination. I therefore conclude that the ecological conditions under which sexual reproduction is favored over asexual reproduction are fairly easily attained and are more general than previous analyses had suggested.     

Loss of complementation and the logic of two-step meiosis

Meiosis is usually a two-step process: two divisions preceded by a duplication. One-step meiosis, a single division without prior replication, is a more logical way to produce haploid gametes; moreover, one-step meiosis leads to higher variabilty in the progeny than two-step meiosis. Yet one-step meiosis is very rare in nature, and may not even exist at all. I suggest that this is because one-step meiosis, in contrast to two-step meiosis, can be easily invaded and replaced by asexual reproduction. I discuss why other existing peculiar forms of division leading to the production of haploid gametes, but not one-step meiosis, have the same effect as two-step meiosis.     

Inverted meiosis and its place in the evolution of sexual reproduction pathways

Inverted meiosis is observed in plants (Cyperaceae and Juncaceae) and insects (Coccoidea, Aphididae) with holocentric chromosomes, the centromeres of which occupy from 70 to 90% of the metaphase chromosome length. In the first meiotic division (meiosis I), chiasmata are formed and rodlike bivalents orient equationally, and in anaphase I, sister chromatids segregate to the poles; the diploid chromosome number is maintained. Non-sister chromatids of homologous chromosomes remain in contact during interkinesis and prophase II and segregate in anaphase II, forming haploid chromosome sets. The segregation of sister chromatids in meiosis I was demonstrated by example of three plant species that were heterozygous for chromosomal rearrangements. In these species, sister chromatids, marked with rearrangement, segregated in anaphase I. Using fluorescent antibodies, it was demonstrated that meiotic recombination enzymes Spo11 and Rad5l, typical of canonical meiosis, functioned at the meiotic prophase I of pollen mother cells of Luzula elegance and Rhynchospora pubera. Moreover, antibodies to synaptonemal complexes proteins ASY1 and ZYP1 were visualized as filamentous structures, pointing to probable formation of synaptonemal complexes. In L. elegance, chiasmata are formed by means of chromatin threads containing satellite DNA. According to the hypothesis of the author of this review, equational division of sister chromatids at meiosis I in the organisms with inverted meiosis can be explained by the absence of specific meiotic proteins (shugoshins). These proteins are able to protect cohesins of holocentric centromeres from hydrolysis by separases at meiosis I, as occurs in the organisms with monocentric chromosomes and canonical meiosis. The basic type of inverted meiosis was described in Coccoidea and Aphididae males. In their females, the variants of parthenogenesis were also observed. Until now, the methods of molecular cytogenetics were not applied for the analysis of inverted meiosis in Coccoidea and Aphididae. Evolutionary, inverted meiosis is thought to have appeared secondarily as an adaptation of the molecular mechanisms of canonical meiosis to chromosome holocentrism.     

DNA repair and the evolution of transformation IV. DNA damage increases transformation

Natural genetic transformation in the bacterium Bacillus subtilis provides a model system to explore the evolutionary function of sexual recombination. In the present work, we study the response of transformation to UV irradiation using donor DNAs that differ in sequence homology to the recipient's chromosome and in the mechanism of transformation. The four donor DNAs used include homologous-chromosomal-DNA, two plasmids containing a fragment of B. subtilis trp⁺ operon DNA and a plasmid with no sequence homology to the recipient cell's DNA. Transformation frequencies for these DNA molecules increase with increasing levels of DNA damage (UV radiation) to recipient cells, only if their transformation requires homologous recombination (i.e. is recA⁺-dependent). Transformation with non-homologous DNA is independent of the recipient's recombination system and transformation frequencies for it do not respond to increases in UV radiation. The transformation frequency for a selectable marker increases in response to DNA damage more dramatically when the locus is present on small, plasmid-borne, homologous fragments than if it is carried on high molecular weight chromosomal fragments. We also study the kinetics of transformation for the different donor DNAs. Different kinetics are observed for homologous transformation depending on whether the homologous locus is carried on a plasmid or on chromosomal fragments. Chromosomal DNA- and non-homologous-plasmid-DNA-mediated transformation is complete (maximal) within several minutes, while transformation with a plasmid containing homologous DNA is still occurring after an hour. The results indicate that DNA damage directly increases rates of homologous recombination and transformation in B. subtilis. The relevance of these results and recent results of other labs to the evolution of transformation are discussed.     

Contrasting patterns of synonymous and nonsynonymous sequence evolution in asexual and sexual freshwater snail lineages

In asexual lineages, both synonymous and nonsynonymous sequence polymorphism may be reduced due to severe founder effects when asexual lineages originate. However, mildly deleterious (nonsynonymous) mutations may accumulate after asexual lineages are formed, because the efficiency of purifying selection is reduced even in the nonrecombining mitochondrial genome. Here we examine patterns of synonymous and nonsynonymous mitochondrial sequence polymorphism in asexual and sexual lineages of the freshwater snail Campeloma. Using clade-specific estimates, we found that synonymous sequence polymorphism was significantly reduced by 75% in asexuals relative to sexuals, whereas nonsynonymous sequence polymorphism did not differ significantly between sexuals and asexuals. Two asexual clades had high negative values for Tajima's D statistic. Coalescent simulations confirmed that various bottleneck scenarios can account for this result. We also used branch-specific estimates of the ratio of amino acid to silent substitutions, K(a)/K(s). Our study revealed that K(a)/K(s) ratios are six times higher in terminal branches of independent asexual lineages compared to sexuals. Coalescent-based reconstruction of gene networks for all sexual and asexual clades indicated that nonsynonymous mutations occurred at a higher frequency in recently derived asexual haplotypes. These findings suggest that patterns of synonymous and nonsynonymous nucleotide polymorphism in asexual snail lineages may be shaped by both severe founder effect and relaxed purifying selection.     

Density-dependent sexual reproduction in natural populations of the rotifer asplanchna girodi

The monogonont rotifer Asplanchna girodi was continuously present in daily and bi-daily plankton samples of Golf Course Pond during the spring of 1977. Two cycles of sexual reproduction occurred during this period. By isolation and culture of females it was possible to determine the reproductive type of collected individuals. Data thus obtained suggest that environmental cues associated with population density are responsible for the production of sexual females.     

Testing some hypotheses on human evolution and sexual dimorphism

Research on human evolution and sexual dimorphism motivates an interesting test problem. In studying hominid phylogeny it is of interest to test whether parallel evolution plays a role. With regard to sexual dimorphism it is of interest to known whether the directions of sexual dimorphism in the populations being compared are the same. We show that testing these two problems gives rise to the same type of hypothesis testing, viz. the problem of testing the hypothesis that the means of independent, normally distributed random vectors with unit covariance matrices are situated on a straight line through the origin. A test is proposed and applied to study the sexual dimorphism of 20 recent skull populations. In this example the hypothesis of equal directions of sexual dimorphism is rejected. The classical theory of constructing multiple discriminant functions (canonical variates) is adapted to the problem of comparing sexual dimorphisms.     

The effects of spatial habitat structure on the evolution of density-dependent growth and reproduction in freshwater snails

We examined the growth and reproductive rates of freshwater snails, Physa acuta, in two habitat types. In the Asabata habitat, snails lived in isolated water pools, which occasionally joined to form a single large pool; in the Kakegawa habitat, they lived in a slow-running water way. Genetic structure assessments using three microsatellite loci supports the idea that a stable panmictic population occupies the Kakegawa habitat. The Asabata habitat, however, is occupied with an alternate mixing population as revealed by microsatellite data. The Asabata population might alternate between localized mating within isolated pools (as revealed by high F _IS and F _IT values) when the water levels are low and panmixia (as revealed by the low F _ST values and AMOVA analysis) when the habitat is flooded. Laboratory experiments, using snails collected from the two habitats, showed that juvenile snails grew faster, laid more eggs, and laid them earlier in the Asabata habitat than in the Kakegawa habitat. Growth rates were lower at high density than at low density in the Kakegawa habitat; the inverse was true in the Asabata habitat. Density-dependent response of individual snail reproduction was higher in the Kakagawa habitat than in the Asabata habitats. The results support the hypothesis that spatial structure affects the evolution of density-dependent growth rates and of timing for reproduction.     

初级代谢产物和有性生殖促进剂对番茄红素发酵的影响

考察几种初级代谢中间产物对三孢布拉霉发酵生产番茄红素的影响,以及卵磷脂对三孢布拉霉正负菌接合孢子及发酵的影响。结果表明：发酵24h分别添加2．0％的柠檬酸和2．0％的三孢酸,番茄红素的产量分别达到0．99g／L和1．26g/L,比对照分别提高39．43％和32．63％;卵磷脂的添加能促进三孢布拉霉两性接合孢子的形成,进而促进番茄红素的合成,当大豆卵磷脂的添加量为0．3％时,番茄红素的产量为1．58g/L,比对照提高56．44％。     

Morphology and evolution of sexual reproduction in the Volvocaceae (Chlorophyta)

Morphological features of sexual reproduction in the Volvocaceae are reviewed, focusing particularly on gametic union and zygote gemination. Both of the two conjugating gametes of the isogamous generaPandorina, Volvulina andYamagishiella bear a tubular mating structure (mating papilla), and plasmogamy is initiated by union of the papillae tips. On zygote germination, a single biflagellate gone cell is released from the zygote wall. Although all the anisogamous and oogamous genera of the Volvocaceae produce “sperm packets” during gametogenesis and a single gone cell at zygote germination, some difference can be recognized in the male gametes. The male gametes ofEudorina bear a tubular cytoplasmic protuberance (putative mating papllla) near the base of the flagella, whereas such a structure recognized at the light microscopic level is not evident inPleodorina andVolvox. Evolution of the sexual reproduction characteristics of volvocacean algae is discussed on the basis of recent cladistic analysis of morphological data as well as of the ribosomal (r) RNA phylogeny and large subunit of the ribulose-1, 5-biphosphate carboxylase/oxygenase(rbcL) gene trees. Dedicated to Emeritus Prof. Hideo Kasaki (Tokyo Metropolitan University) on the occasion of his 77th birthday. Recipient of the Botanical Society Award for Young Scientists, 1994.     

Optimal allocation to vegetative and sexual reproduction in plants: the effect of ramet density

Vegetative reproduction is a very common alternative by which plants can contribute to the next generations. There are many considerations predicting which mode of reproduction, vegetative or sexual, should be favored and numerous experimental studies to verify them. However, the results are inconsistent especially when the effect of plant density is considered. I apply here a dynamic optimization model to predict the rate of vegetative and sexual reproduction in plants as a response to changes in the local plant density. The population is assumed to occupy a heterogeneous environment consisting of patches in which growth and reproduction of plants are possible and unfavorable space between them. As the environment is globally stable, the seeds, which can disperse without restriction, exhibit a constant recruitment rate. The ramets are assumed to settle only within the patch of the mother plant. The rate of ramet production effects local density, which in turn determines ramet recruitment. The optimal strategy maximizes the expected lifetime genetic contribution, realized via both vegetative and sexual reproduction. The solutions obtained under these assumptions are dualistic. The model predicts that different approaches applied in studying the effect of ramet density should give opposite outcomes. When the comparison is between patches in natural populations, a positive relationship between relative ramet allocation and density is expected. When the density is experimentally manipulated or its effect is analyzed across different successional stages, a negative relationship should be found. The results seem to be confirmed by empirical studies.     

The pattern of population growth as a function of redundancy and repair

A basic model of hierarchical structure, expressed by simple, linear differential equations, shows that the pattern of population growth is essentially determined by conditions of redundancy in the sub-structure of individuals. There does not exist any possible combination between growth rate and accident rate that could balance population numbers and/or the level of redundancy within the population; all possible combinations either lead to extinction or to positive population growth with a decline of the fraction of individuals with redundant substructure. Declining populations, however, can be held fluctuating between certain limits by periodic phases of sub-unit repair. These results are particularly pertinent to the population dynamics of diploid (polyploid) organisms.