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Outcrossing rate and inbreeding depression in the perennial yellow bush lupine, Lupinus arboreus (Fabaceae).

Little is known about the breeding systems of perennial Lupinus species. We provide information about the breeding system of the perennial yellow bush lupine, Lupinus arboreus, specifically determining self-compatibility, outcrossing rate, and level of inbreeding depression. Flowers are self-compatible, but autonomous self-fertilization rarely occurs; thus selfed seed are a product of facilitated selfing. Based on four isozyme loci from 34 maternal progeny arrays of seeds we estimated an outcrossing rate of 0.78. However, when we accounted for differential maturation of selfed seeds, the outcrossing rate at fertilization was lower, ∼0.64. Fitness and inbreeding depression of 11 selfed and outcrossed families were measured at four stages: seed maturation, seedling emergence, seedling survivorship, and growth at 12 wk. Cumulative inbreeding depression across all four life stages averaged 0.59, although variation existed between families for the magnitude of inbreeding depression. Inbreeding depression was not manifest uniformly across all four life stages. Outcrossed flowers produced twice as many seeds as selfed flowers, but the mean performance of selfed and outcrossed progeny was not different for emergence, seedling survivorship, and size at 12 wk. Counter to assumptions about this species, L. arboreus is both self-compatible and outcrosses ∼78% of the time.

Journal Article↗

Cleistogamous capitulum in Centaurea melitensis (Asteraceae): heterochronic origin.

Cleistogamous capitula formed by Centaurea melitensis display a number of morphological and functional changes with respect to chasmogamous capitula that ensure self-fertilization. Because no studies have hitherto addressed the evolution of cleistogamy in Asteraceae, it was considered useful to ascertain whether these changes are attributable to one or more of the heterochronic processes reported in the literature. Bivariate allometric analyses were performed, and changes were represented graphically using Gould's clock models for size, shape, and age of several capitulum and floret structures. Results suggest that the partially paedomorphic appearance of cleistogamous with respect to chasmogamous capitula is attributable to three processes: (1) early onset of floral development (predisplacement), (2) decreased growth rate of the whorls studied (except gynoecium width) and (3) early offset time (progenesis). The latter appears to play the most significant role in the origin of the cleistogamous capitulum.

Journal Article↗

Transposon dynamics and the breeding system.

The selfish DNA hypothesis predicts that natural selection is responsible for preventing the unregulated build up of transposable elements in organismal genomes. Accordingly, between-species differences in the strength and effectiveness of selection against transposons should be important in driving the evolution of transposon activity and abundance. We used a modeling approach to investigate how the rate of self-fertilization influences the population dynamics of transposable elements. Contrasting effects of the breeding system were observed under selection based on transposon disruption of gene function versus selection based on element-mediated ectopic exchange. This suggests that the comparison of TE copy number in organisms with different breeding systems may provide a test of the relative importance of these forces in regulating transposon multiplication. The effects of breeding system also interacted with population size, particularly when there was no element excision. The strength and effectiveness of selection against transposons was reflected not only in their equilibrium abundance, but also in the per-site element frequency of individual insertions and the coefficient of variation in copy number. These results are discussed in relation to evidence on transposon abundance available from the literature, and suggestions for future data collection.

Breeding↗

Experimental analysis of protogyny in Aquilegia canadensis (Ranunculaceae).

Dichogamy is very common in flowering plants and is widely thought to reduce pollen-pistil interference, especially self-pollination. Yet, the functional significance of dichogamy has rarely been studied. We investigated the nature and functioning of dichogamy in eastern Ontario populations of Aquilegia canadensis, a highly selfing columbine previously described as protogynous. We then manipulated flowers to determine whether increased protogyny would reduce self-fertilization. Contrary to previous reports, A. canadensis is not dichogamous. Controlled pollinations in a greenhouse showed that pollen tubes generally begin to develop after anther dehiscence. Although stigmas can collect pollen early in floral development, naturally pollinated flowers collected from four populations had few pollen grains on stigmas and almost no pollen tubes in styles until after anther dehiscence. Limited pollen deposition before anther dehiscence was also associated with low nectar availability and limited sepal expansion. Because inbreeding depression is strong in this species, selection may favor increased protogyny if it reduces selfing. We tested this hypothesis by comparing the level of selfing in flowers rendered protogynous by the removal of the first 19 (of 39) anthers to develop, with nonprotogynous control flowers. Contrary to expectations, protogyny did not reduce selfing. Our results emphasize the importance of detailed field observations and manipulative experiments in understanding the nature and functional significance of dichogamy.

Journal Article↗

Evolution of the self-pollinating flower in Clarkia xantiana (Onagraceae). I. Size and development of floral organs.

Clarkia xantiana has two subspecies that differ in breeding system: ssp. xantiana, which is outcrossing, and ssp. parviflora, which is self-fertilizing. Outcrossing is the ancestral breeding system for the genus Clarkia. Flowers of ssp. parviflora have characteristics commonly associated with selfing taxa: they are smaller and have little temporal and spatial separation between mature anthers and stigma (dichogamy and herkogamy, respectively). Flower morphology and development were studied in four populations of each subspecies to establish the developmental changes that occurred in the evolution of selfing. In particular, we sought to evaluate the hypothesis that the selfing flower may have arisen as a byproduct of selection for rapid maturation in the arid environment occupied by ssp. parviflora. This hypothesis predicts that development time should be reduced in spp. parviflora relative to ssp. xantiana. We also sought to compare the pattern of covariation of flower morphology and development between subspecies to that within subspecies. Similar within vs. between patterns of covariation could be indicative of developmental or functional constraints on the independent evolution of floral parts. In spite of significant variation among populations within subspecies, the subspecies clearly differ in flower morphology and development. All floral organs, except ovaries, are smaller in ssp. parviflora than in ssp. xantiana. The flower plastochron, the duration of flower development from bud initiation to anthesis, and the duration of protandry are all shorter in ssp. parviflora than in ssp. xantiana. Maximum relative growth rates are higher for all organs in ssp. parviflora than in ssp. xantiana. Thus, progenesis (i.e., via a reduction in development time) is combined with growth acceleration in the evolution of the selfing flower. Since reduced development time and growth acceleration both allow selfing flowers to mature earlier than outcrossing ones, selection for early maturation may have contributed to the evolution of the selfing flower form. The pattern of trait covariation differs within spp. parviflora relative to the patterns within spp. xantiana and between the two subspecies, suggesting that floral parts can and have evolved independently of one another.

Journal Article↗

The molecular basis of the high linoleic acid content in Petunia seed oil: analysis of a seed-specific linoleic acid mutant.

From a random transposon mutagenesis experiment, using Petunia line W138, a seed-specific linoleic acid mutant was isolated. The tagged gene was cloned and identified as a microsomal Delta(12) desaturase. Expression of the gene, however, was constitutive and not, as might have been expected, seed-specific. Moreover, self-fertilized homozygous mutants still contain 40% 18:2 in the seed lipid fraction. This suggests that at least two (seed-specific) Delta(12) desaturase genes are responsible for the high linoleic acid content in Petunia seed oil. Five members of the microsomal Delta(12) desaturase gene family have been identified and isolated. Data are presented on the molecular characterization and tissue-specific expression of these genes, which suggest that, in Petunia, the flux through the prokaryotic and eukaryotic pathways of lipid synthesis might be different from the situation found in Arabidopsis.

Cloning, Molecular↗

[Genetic analysis of T1 progeny of transgenic tobacco with metallothionein gene and metallothionein domain mutant alpha alpha gene].

The chimeric gene containing a cloned mouse metallothionein processed gene or a cloned mouse metallothionein domain mutant alpha alpha gene was respectively introduced into tobacco (Nicotiana tobacum L. cv. NC89) on a disarmed Ti-plasmid of Agrobacterium tumefaciens. T1 seeds from self-fertilized transgenic tobacco were germinated on media containing cadmium or herbicide PPT. The PPT tolerance trait followed Mendelian inheritance and co-segregated with heavy metal tolerance. Meanwhile Southern blot and Western blot verified the existence of the MT gene and alpha alpha mutant gene in the T1 generation plants which keep tolerance to heavy metal. All the results demonstrated the stable integration and inheritance of exotic genes. In addition, assay of the root length and fresh weight of T1 seedlings indicate that transgenic tobacco plants with alpha alpha mutant gene still have a little higher tolerance than that with matural MT gene.

Animals↗

Brachypodium distachyon. A new model system for functional genomics in grasses.

A new model for grass functional genomics is described based on Brachypodium distachyon, which in the evolution of the Pooideae diverged just prior to the clade of "core pooid" genera that contain the majority of important temperate cereals and forage grasses. Diploid ecotypes of B. distachyon (2n = 10) have five easily distinguishable chromosomes that display high levels of chiasma formation at meiosis. The B. distachyon nuclear genome was indistinguishable in size from that of Arabidopsis, making it the simplest genome described in grasses to date. B. distachyon is a self-fertile, inbreeding annual with a life cycle of less than 4 months. These features, coupled with its small size (approximately 20 cm at maturity), lack of seed-head shatter, and undemanding growth requirements should make it amenable to high-throughput genetics and mutant screens. Immature embryos exhibited a high capacity for plant regeneration via somatic embryogenesis. Regenerated plants display very low levels of albinism and have normal fertility. A simple transformation system has been developed based on microprojectile bombardment of embryogenic callus and hygromycin selection. Selected B. distachyon ecotypes were resistant to all tested cereal-adapted Blumeria graminis species and cereal brown rusts (Puccinia reconditia). In contrast, different ecotypes displayed resistance or disease symptoms following challenge with the rice blast pathogen (Magnaporthe grisea) and wheat/barley yellow stripe rusts (Puccinia striformis). Despite its small stature, B. distachyon has large seeds that should prove useful for studies on grain filling. Such biological characteristics represent important traits for study in temperate cereals.

Anti-Bacterial Agents↗

[Biological characteristics of different forest soils in Nanjing-Zhenjiang mountan area].

Studies on the biological characteristics, including soil microbe, soil enzyme activity, soil nutritient content, and litter decomposition of different forest soils in Nanjing-Zhenjiang mountain area showed that the amounts of microbes and the activities of six enzymes in forest soils changed regularly in different forests during different months. The contents of nutritional elements varied regularly with forest growth bio-cycles, and were inerrelated prominently with the amounts of soil microbes and the activities of soil enzymes. There existed temporal-spatial differences in the decomposition rate of litters and the reverted velocity of nutrients among different forests. The comparisons of various biological characteristics among secondary Quercus variabilis forest, Phllostachys pubescens forest, and Cunninghamia lanceolata forest indicated that secondary Quercus variabilis forest had the most abundant nutrients in soil, and possessed the strongest ability of self-fertilization. Therefore, to construct coniferous forests with broadleaf trees in this area could avoid or abate the decline of soil fertility.

Biomass↗

Major transitions in evolution by genome fusions: from prokaryotes to eukaryotes, metazoans, bilaterians and vertebrates.

The major transitions in human evolution from prokaryotes to eukaryotes, from protozoans to metazoans, from the first animals to bilaterians and finally from a primitive chordate to vertebrates were all accompanied by increases in genome complexity. Rare fusion of divergent genomes rather than continuous single gene duplications could explain these jumps in evolution. The origin of eukaryotes was proposed to be due to a symbiosis of Archaea and Bacteria. Symbiosis is clearly seen as the source for mitochondria. A fundamental difference of higher eukaryotes is the cycle from haploidy to diploidy, a well-regulated genome duplication. Of course, self-fertilization exists, but the potential of sex increases with the difference of the haploid stages, such as the sperm and the egg. What should be the advantage of having two identical copies of a gene? Still, genes duplicate all the time and even genomes duplicate rather often. In plants, polyploidy is well recognized, but seems to be abundant in fungi and even in animals, too. However, hybridization, rather than autopolyploidy, seems to be the potential mechanism for creating something new. The problem with chimaeric, symbiotic or reticulate evolution events is that they blur phylogenetic lineages. Unrecognized paralogous genes or random loss of one of the paralogs in different lineages can lead to false conclusions. Horizontal genome transfer, genome fusion or hybridization might be only truly innovative combined with rare geological transitions such as change to an oxygen atmosphere, snowball Earth events or the Cambrian explosion, but correlates well with the major transitions in evolution.

Animals↗

Mating behavior in the pulmonate small melampus: can regeneration restore function?

Previous anatomical observations have established that the penial complex of adult Melampus bidentatus is specifically reinnervated and that the penial complex itself can regenerate. This review describes experiments to determine whether a reinnervated penial complex and a regenerated penial complex can function in mating. The snails are not self-fertile, so the ability to transfer sperm, evaluated by production of fertile eggs, was the test of successful regeneration. Snails with nerve transections produced fertile eggs within the time-span expected for axonal regeneration to the target organ. Penial complex ablation interfered with sperm transfer for two egg-laying cycles but fertile eggs were laid within a time-span consistent with regeneration of a penial complex.

Animals↗

Evolutionary implications of a high selfing rate in the freshwater snail Lymnaea truncatula.

Self-compatible hermaphroditic organisms that mix self-fertilization and outcrossing are of great interest for investigating the evolution of mating systems. We investigate the evolution of selfing in Lymnaea truncatula, a self-compatible hermaphroditic freshwater snail. We first analyze the consequences of selfing in terms of genetic variability within and among populations and then investigate how these consequences along with the species ecology (harshness of the habitat and parasitism) might govern the evolution of selfing. Snails from 13 localities (classified as temporary or permanent depending on their water availability) were sampled in western Switzerland and genotyped for seven microsatellite loci. F(IS) (estimated on adults) and progeny array analyses (on hatchlings) provided similar selfing rate estimates of 80%. Populations presented a low polymorphism and were highly differentiated (F(ST) = 0.58). Although the reproductive assurance hypothesis would predict higher selfing rate in temporary populations, no difference in selfing level was observed between temporary and permanent populations. However, allelic richness and gene diversity declined in temporary habitats, presumably reflecting drift. Infection levels varied but were not simply related to either estimated population selfing rate or to differences in heterozygosity. These findings and the similar selfing rates estimated for hatchlings and adults suggest that within-population inbreeding depression is low in L. truncatula.

Alleles↗

[Mechanisms of meiotic restitution and their genetic regulation in wheat-rye polyhaploids].

Meiosis has been studied in partially fertile wheat-rye F1 hybrids yielded by crosses Triticum aestivum (Saratovskaya 29 variety) x Secale cereale L. (Onokhoiskaya variety) (4x = 28). Hybrid self-fertility proved to be caused by formation of restituted nuclei, which appear after equational segregation of univalent chromosome in AI and sister chromatid non-separation in AII of meiosis, as well as after AI blockage in three different ways. Both types of meiotic restitution were found in each hybrid plant. Expression of the "meiotic restitution" trait varied significantly in polyhaploids of the same genotype (ears of the same plants, anthers of the same ear, microsporocytes of the same anther). Chromatin condensation in prophase proved to be related to the division type and univalent segregation in AI. During reduction segregation of univalents in AI, sister chromatid cohesion and chromosome supercondensation remained unchanged. The results obtained suggest that in the remote hybrids with haploid karyotype of the parental origin (polyhaploids), the program of two-stage meiosis may be fundamentally transformed to ensure one instead of two divisions. We propose that meiotic restitution is a result of special genetic regulation of the kinetochore organization (both structural and functional) and chromatin condensation, i.e. of major meiotic mechanisms.

Haploidy↗

[Identification of the apomixis in Poa pratensis L. using SDS-electrophoresis of endosperm reserve proteins].

To determine the characteristics of variability of seed reproduction in the Kentucky bluegrass (Poa pratensis), individual seed variability with respect to the composition of endospermal reserve proteins was studied. Comparative analysis of caryopses obtained by self-fertilization and free fertilization of plants I1 (no. 2-4) and I2 (no. 2-4-7) of the wild-type specimen Murmanskii-95 was performed using SDS-PAGE. Using a cytoembryological express method, we demonstrated that facultative stimulation-autonomous apomeiotic apomixis, along with the formation of meiotic megasporocytes, is characteristic of the Kentucky bluegrass. This method made it possible to determine the consequences of meiotic processes in the maternal plant and to reveal the hybrid nature of seed endosperm.

Crosses, Genetic↗

Sperm-limited fecundity in nematodes: how many sperm are enough?

The Bateman principle, which holds that oocytes are the limiting gamete in reproduction, is violated in a variety of species. Self-fertilizing hermaphrodites of the nematode Caenorhabditis elegans provide an example of a system in which sperm number limits lifetime reproductive output, in this species due to the protandrous nature of sperm production that in turn delays the onset of fertilization. This reproductive delay forms the basis of a trade-off between generation time and total fecundity, in which sperm number plays a pivotal role. I use an age-structured population model to describe the number of sperm that maximize fitness, given larval development time and rates of gamete production. The model predicts the evolution of sperm numbers that are consistent with empirical data for C. elegans provided that precocious larval sperm production is taken into account. Several testable hypotheses follow from the model regarding how natural selection and environmental variation may influence patterns of sperm production among populations or species with a similar mode of reproduction.

Age Factors↗

[Intraspecific structure and reproductive relationships between Elymus mutabilis and E. transbaicalensis (Poaceae) in southern Siberia from the viewpoint of taxonomical genetics].

The relationships between StH-genomic species Elymus mutabilis (Drob.) Tzvel. and E. transbaicalensis (Nevski) Tzvel. were examined using biosystematic and biochemical (SDS electrophoresis of endosperm proteins) methods. The results showed the following. (1) There is interspecific morphological discreteness that permits assigning most individuals from the wild to a definite species. However, self-fertile intermediate individuals of introgressive origin were found. (2) Both taxa exhibited marked specificity of electrophoretic characteristics of endosperm proteins. (3) The species form two separate recombination gene pools (RGPs). Thus, E. mutabilis and E. transbaicalensis should be regarded as two distinct species. Nevertheless, there is some genetic exchange and unidirectional gene flow between the RGPs, especially in secondary (successive) habitats where mixed populations are observed. Two RGPs form the single introgressive gene pool (IGP), which includes other StH-genomic species of Northern Eurasia.

Electrophoresis, Polyacrylamide Gel↗

[Morphology, cytogenetics of intergeneric hybrid between Aegilops taucchii and Dasypyrum villosum].

"Aegilops tauschii x Dasypyrum villosum" F1 hybrids were obtained by the combination of hybridization and embryo culture in vitro. Chromosome pairing behavior in meiosis of the hybrid F1 was carried out. Results showed that in an average , 1.25 rod bivalents were observed in one PMC, meiotic configuration was 2n=14=11.49 I + 1.25 II (Xta=1.25) and most of PMCs possessed 1 approximately 5(rod) bivalens, indicating that the relatively high homeology was detected between the D genome of Ae. tauschii and the V genome of D. villosum. The morphological differences between F1 hybrids and their parents were significant. F1 plants were highly self-sterile, but partially self-fertile after treated by chromosome doubling technique.

Chimera↗

Cytology and fertility of viable hybrids of Nicotiana tabacum L. cv. TB-566 with N. alata Link et Otto.

A vigorous hybrid (N. tabacum cv. TB-566 tetra x N. alata) x N. alata was obtained by backcrossing a partly viable sesquidiploid hybrid N. tabacum cv. TB-566 tetra x N. alata to N. alata. The hybrid was a 35-chromosome near-amphihaploid with a pair of N. alata chromosomes in disomic condition. It was completely self- and cross-sterile and formed from 7 to 8 bivalents in pollen mother cells. By using stem pith culture, polyploidized regenerants were obtained from the 35-chromosome hybrid with somatic chromosome numbers from 65 to 70. These regenerants showed fairly regular meiosis with the number of bivalents in pollen mother cells ranging from 27.3 to 30.4. Irregularities in meiosis included a high number of univalents, aberrant tetrads, and a high frequency of micronuclei. The percentage of acetocarmine-stainable pollen ranged from 22.1 to 78.4. A 66-chromosome regenerant showed fairly regular meiosis and was self-fertile but could not be backcrossed to N. tabacum. This barrier seems to be caused by genic imbalance rather than irregularities of meiotic divisions. Hence transfer programs based on the introgression of entire linkage groups (sexual and somatic hybridization) seem to be of little use in the case of that species.

Chromosomes, Plant↗