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Autotoxicity in Lolium rigidum: analyzing the role of chemically mediated interactions in annual plant populations.

Autotoxicity is the chemical influence of one plant on the development of individuals of the same species. Autotoxicity in perennial plants can be a useful mechanism to avoid future conspecific competitors, however, the ecological role of autotoxicity in annuals, if any, remains uncertain. In this paper, we analyse the autotoxic phenomenon in Lolium rigidum, an annual, self-incompatible grass from semi-arid environments suspected to be autotoxic, and we dissert on the mechanisms of intraspecific interference, carrying out density-dependent experiments with additions of conspecific competitors and plant residues. Our experiments show that chemical interference and resource competition are density-dependent processes that act in opposite directions. Inhibitory chemical interference increases at low population densities whereas competition for resources intensifies at high densities. Based on these results, the manuscript theorizes on the ecological role of the autotoxic phenomenon in short-lived plants. We hypothesize that autotoxicity enhances population viability by restraining the development of populations with few individuals, which should decrease the chances of outcross and experience inbreeding depression. If this hypothesis proves true, it could contribute to explain the unusually high within-population diversity encountered in this species, which contrasts with reported values from other similar wind-pollinated, self-incompatible grasses.

Lolium↗

Plants, pairing and phenotypes--two's company?

An RNA-based communication network appears to play a crucial role in regulating gene expression and in repressing viral and transposon sequences in plant genomes. In this article, we consider the evidence that gene expression might also be controlled epigenetically at a level other than non-coding RNA species-chromosome pairing. This epigenetic communication between sequences might be based--as it is in other organisms--on the physical pairing between homologues and the transfer of information between corresponding epigenetic landscapes. We suggest that paramutation might represent just one--albeit extreme and obvious--facet of a pairing-based gene expression regulation system in plants. Further exciting evidence for pairing occurring between homologues in plants is now mounting. An appreciation that pairing interactions might be important throughout plant development could assist in understanding phenomena such as endosperm imprinting, hybrid phenotypes and inbreeding depression.

Alleles↗

The genetic architecture of house fly mating behavior.

This chapter summarizes several experimental approaches used to identify the effects of dominance, epistasis, and genotype-by-environment interactions in the genetic architecture of the mating behavior of the common house fly (Musca domestica L.). Quantitative genetic investigations of mating behavior hold special intrigue for unraveling the complexities of fitness traits, with applications to theory on sexual selection and speciation. Besides being well suited to large-scale quantitative genetic protocols, the house fly has a remarkably complex courtship repertoire, affording special opportunities for studies on communication, social interactions, and learning. Increased additive genetic variances for the courtship repertoire of experimentally bottlenecked populations provided evidence for the presence of dominance and/or epistasis. Negative genetic variances in these populations suggested genotype-by-environment interactions, where the environment is the mating partner. Line cross assays of populations that had been subjected to selection for divergent courtship repertoire confirmed that both dominance and epistasis have significant effects. These crosses also showed more directly that the expression of the male's genotype is dependent upon the preferences of his mating partner. Repeatability studies also detailed how males alter their courtship performances with successive encounters within and across females, such that the males learn to improve their techniques in securing copulations. A review of 41 animal behavior studies found that a wide range of traits and taxa have dominance, epistasis, and genotype-y-environment interactions, although house fly courtship may remain a unique model where learning is an intersexually selected trait. Future development of more sophisticated molecular techniques for the M. domestica genome will help unravel the underlying biochemical and developmental pathways of these quantitative genetic interactions for a more complete understanding of the processes of inbreeding depression, outbreeding depression, and pleiotropy.

Animals↗

Restoration of genetic variation lost - the genetic rescue hypothesis.

It has been long known that immigrants from surrounding populations might prevent the extinction of small populations, a process known as the 'rescue effect'. This focuses on the demographic effects of migration through the direct positive influence that immigrants have on abundance of the recipient population. Now, two recent papers have indicated another potentially important way that migration might rescue populations from extinction - replenishing genetic variation and reducing inbreeding depression, or what has been termed 'genetic rescue'.

Journal Article↗

Phenotypic plasticity for life-history traits in Drosophila melanogaster. III. Effect of the environment on genetic parameters.

We estimated genetic and environmental variance components for developmental time and dry weight at eclosion in Drosophila melanogaster raised in ten different environments (all combinations of 22, 25 and 28 degrees C and 0.5, 1 and 4% yeast concentration, and 0.25% yeast at 25 degrees C). We used six homozygous lines derived from a natural population for complete diallel crosses in each environment. Additive genetic variances were consistently low for both traits (h2 around 10%). The additive genetic variance of developmental time was larger at lower yeast concentrations, but the heritability did not increase because other components were also larger. The additive genetic effects of the six parental lines changed ranks across environments, suggesting a mechanism for the maintenance of genetic variation in heterogenous environments. The variance due to non-directional dominance was small in most environments. However, there was directional dominance in the form of inbreeding depression for both traits. It was pronounced at high yeast levels and temperatures but disappeared when yeast or temperature were decreased. This meant that the heterozygous flies were more sensitive to environmental differences than homozygous flies. Because dominance effects are not heritable, this suggests that the evolution of plasticity can be constrained when dominance effects are important as a mechanism for plasticity.

Adaptation, Physiological↗

Brother-sister marriage in Roman Egypt.

According to official census returns from Roman Egypt (first to third centuries CE) preserved on papyrus, 23.5% of all documented marriages in the Arsinoites district in the Fayum (n = 102) were between brothers and sisters. In the second century CE, the rates were 37% in the city of Arsinoe and 18.9% in the surrounding villages. Documented pedigrees suggest a minimum mean level of inbreeding equivalent to a coefficient of inbreeding of 0.0975 in second century CE Arsinoe. Undocumented sources of inbreeding and an estimate based on the frequency of close-kin unions (corrected downwards to 30% for Arsinoe) indicate a mean coefficient of inbreeding of F = 0.15-0.20 in Arsinoe and of F = 0.10-0.15 in the villages at the end of the second century CE. These values are several times as high as any other documented levels of inbreeding. A schematic estimate of inbreeding depression in the offspring of full sibling couples indicates that fertility in these families had to be 20-50% above average to attain reproduction at replacement level. In the absence of information on the amount of genetic load in this population, this estimate may be too high.

Censuses↗

Evidence for strategic egg production in a hermaphroditic cestode.

The cestode Schistocephalus solidus is a simultaneous hermaphrodite that grows in 2 intermediate hosts and reproduces rapidly within a few days in the gut of a bird. Reproduction takes place by self- or cross-fertilization. Here, it was tested whether egg production differs between S. solidus that reproduce alone and those that are allowed to reproduce in pairs. Egg production in an in vitro system was found to depend on the cestodes' social situation. When kept alone, larger cestodes produced larger eggs. This was not so when kept in pairs--the difference between these 2 reproductive modes being highly significant in this respect. Further experiments revealed that, within the first 3 days, these hermaphrodites produced a larger total egg mass when kept alone than when kept in pairs. This was also reflected by the energy contents of the cestodes after this time-span: selfers had used up more energy than paired worms. Furthermore, S. solidus appeared to adjust its investment per egg depending on whether the offspring will be the result of self- or cross-fertilization. Selfers produced larger numbers of eggs, but these eggs were smaller and contained even smaller embryos per given egg size than eggs of potentially outbreeding cestodes. Selfed eggs reached lower hatching rates. Although this is to be expected from inbreeding depression it may also be an effect of the reduced maternal investment per egg. The observed phenotypic plasticity in the reproduction of S. solidus is discussed within 4 evolutionary frameworks: local mate competition adjusted for hermaphrodites, the hermaphrodite's dilemma, bet-hedging, and sib-competition.

Analysis of Variance↗

Mitochondrial DNA sequence divergence among some west European brown bear (Ursus arctos L.) populations. Lessons for conservation.

We used the polymerase chain reaction and nucleotide sequencing to study mitochondrial DNA (mtDNA) genetic variability of brown bears (Ursus arctos) belonging to some western European populations. Demographic decline and isolation have apparently shaped genetic variability within and between populations. Small relict bear populations were apparently monomorphic and fixed for different mtDNA haplotypes. Genetic relationships among mtDNA haplotypes suggested the existence of a phylogeographic structure. We discuss the possible roles of historical and demographic factors in determining the observed genetic patterns. The low genetically effective size of the studied brown bear populations points to risks of continuing future loss of gene diversity and increase in inbreeding. Artificial gene flow through restocking could alleviate inbreeding depression in small isolated brown bear populations, and mtDNA phylogeographic relationships may provide a framework for their genetic management.

Animals↗

Genetic substructuring as a result of barriers to gene flow in urban Rana temporaria (common frog) populations: implications for biodiversity conservation.

The ability to maintain small populations in quasi-natural settings is an issue of considerable importance in biodiversity conservation. The genetic structure of urban common frog (Rana temporaria) populations was determined by allozyme electrophoresis and used to evaluate the effects of restricted intersite migration. Despite the lack of any absolute barrier to movement between ponds, substantial genetic differentiation was found between sites separated by an average of only 2.3 km. Genetic distances between these town ponds correlated positively with geographical distances and were almost twice as great as those found between rural sites separated by an average of 41 km. Measures of genetic diversity and fitness were always lowest in the town, where the degree of subpopulation differentiation (FST = 0.388) was high. Population decline was not evident in the town, but molecular and fitness data indicated the presence of genetic drift and inbreeding depression. The long-term survival of artificially restricted populations, particularly of relatively sedentary species, may require molecular monitoring, if genetic diversity is not to be lost by chance when facets of the species niche prove to be poorly understood.

Alleles↗

Context-dependent autonomous self-fertilization yields reproductive assurance and mixed mating.

The evolution of self-fertilization in hermaphrodites is opposed by costs that decrease the value of self progeny relative to that of outcross progeny. However, self-fertilization is common in plants; 20% are highly selfing and 33% are intermediate between selfing and outcrossing. Darwin proposed an adaptive benefit of self-pollination in providing reproductive assurance when outcrossing is impossible. Moreover, if outcross pollen receipt is inconsistent within or between years, these conditions likewise favour self-pollination, and this can result in a mixture of self and outcross seed production (mixed mating). Despite wide acceptance, the reproductive assurance hypothesis has lacked the support of complete empirical evidence to show that variable pollination can create both the ecological and genetic conditions favouring self-pollination. We recently showed in Collinsia verna that during periods of infrequent pollinator visits, autonomous self-pollination boosted seed output per flower, the key ecological condition. Here we show low inbreeding depression and marker-based estimates of selfing, demonstrating that when the pollination environment in wild populations necessitates reproductive assurance, selfing rates increase. We provide a complete demonstration of reproductive assurance under variable pollination environments and mechanistically link reproductive assurance to intermediate selfing rates through mixed mating.

Breeding↗

Developmental stability in Brassica cretica: the effect of crossing distance on fluctuating asymmetry in cotyledon morphology.

In the present investigation of Brassica cretica, a wild relative of the cultivated cabbage, B. oleracea, we performed an extensive crossing experiment, involving self-pollinations, random outcrosses within populations and hybridizations between populations or species, to evaluate the relationship between crossing distance and developmental stability, estimated as the absolute difference between the right and left lobe of the cotyledons. The frequency distribution of the right-minus-left scores had a narrower peak than expected for normally-distributed data, but there was no directional asymmetry or antisymmetry. Despite evidence for inbreeding depression in seedling biomass and cotyledon size, the type of cross had negligible influence on cotyledon asymmetry. Separate analyses of between-population hybrids revealed differences among progenies from different pairs of populations and a tendency for the F1 hybrid means to decrease with the geographic distance separating the parent populations, but only for the two size variables. Based on these and other observations, we propose that the degree of cotyledon asymmetry is unrelated to genome-wide characteristics, such as the level of heterozygosity and genomic co-adaptation, and that cotyledon asymmetry is unrelated to the level of genetic stress experienced by each individual. Hence, there is no reason to consider measures of asymmetry as more sensitive indicators of genetic health than conventional fitness variables.

Analysis of Variance↗

Mating system and population genetic structure of an androdioecious tree, Fraxinus lanuginosa Koidz. (Oleaceae) in northern Japan.

Models for the maintenance of androdioecy have suggested that selfing of hermaphrodites decreases the frequency of males in a population (the "male frequency"). To test this hypothesis, we used electrophoretic allozyme methods to study the mating system and population genetics of an androdioecious tree, Fraxinus lanuginosa, which exhibits large variations in male frequency among subpopulations in central Hokkaido (northern Japan). We estimated the outcrossing rates by using seeds assayed at three polymorphic loci, and found that the multilocus outcrossing rate (t(m)) increased with increasing male frequency (q) (t(m) = 0.69 to 0.99, q = 0.11 to 0.59). Fixation indices (F(j)) estimated from these t(m) values ranged from 0.006 to 0.185, and were significantly greater than zero in plots with a low male frequency (q or = 0.47). However, fixation indices at the late life stage (F(a)) were not significantly different from zero at five loci (F(a) = 0.056 to 0.101, q = 0.11 to 0.61), and the F(a) values were lower than the F(j) values in several of the plots where both values were measured. These results indicate that inbreeding depression substantially decreases the proportion of selfed progeny in partially self-fertilising subpopulations. A theoretical model for the maintenance of androdioecy showed expected male frequencies significantly lower than the observed values in plots with high male frequency (q = 0.59 to 0.61), although the differences between the expected and observed values of male frequencies were not significant in plots with intermediate or low male-frequencies (q = 0.11 to 0.47). In this paper, the factors are discussed that affect variations in the male frequency among subpopulations of F. lanuginosa based on these findings.

Animals↗

Genetic and demographic dynamics of small populations of Silene latifolia.

Small local populations of Silene alba, a short-lived herbaceous plant, were sampled in 1994 and again in 1999. Sampling included estimates of population size and genetic diversity, as measured at six polymorphic allozyme loci. When averaged across populations, there was very little change between samples (about three generations) in population size, measures of within-population genetic diversity such as number of alleles or expected heterozygosity, or in the apportionment of genetic diversity within and among populations as measured by F(st). However, individual populations changed considerably, both in terms of numbers of individuals and genetic composition. Some populations doubled in size between samples, while others shrank by more than 75%. Similarly, expected heterozygosity and allele number increased by more than two-fold in individual populations and decreased by more than three-fold in others. When population-specific change in number and change in measures of genetic diversity were considered together, significant positive correlations were found between the demographic and genetic variables. It is speculated that some populations were released from the demographic consequences of inbreeding depression by gene flow.

Alleles↗

Molecular differentiation within and among island populations of the endemic plant Scalesia affinis (Asteraceae) from the Galápagos Islands.

Molecular variance was estimated in seven populations of the endemic species Scalesia affinis within and among islands of the Galapagos. The analysis, based on 157 polymorphic AFLP markers, revealed a high differentiation among populations, of which most was partitioned among islands. In addition, the information content of AFLP markers was tested with sets of discriminant analyses based on different numbers of AFLP markers. This indicated that the markers were highly informative in discriminating the populations. Although one of four populations from the island Isabela was sampled from a volcano 100 km away from the remaining populations, this population resembled the others on Isabela. The partitioning of molecular variance (AFLP) resulted in two unities, one consisting of populations from Isabela and one of populations from Santa Cruz and Floreana. The differentiation in two chloroplast microsatellites was higher than for AFLP markers and equally partitioned among populations within islands as among islands. Thus, gene flow via fruits within islands is as limited as among islands. The lower differentiation within islands in the nuclear AFLP markers may thus indicate that gene flow within islands is mostly accounted for by pollen transfer. S. affinis is the only species in the genus that is not listed in 2000 IUCN Red List of Threatened Species. However, due to prominent grazing and land exploitation, some populations have recently been reduced markedly, which was reflected in lower diversity. As inbreeding depression is present in the species, the rapid bottlenecks are threats to the populations.

Asteraceae↗

To self, or not to self... a review of outcrossing and pollen-mediated gene flow in neotropical trees.

Despite the typically low population densities and animal-mediated pollination of tropical forest trees, outcrossing and long-distance pollen dispersal are the norm. We reviewed the genetic literature on mating systems and pollen dispersal for neotropical trees to identify the ecological and phylogenetic correlates. The 36 studies surveyed found >90% outcrossed mating for 45 hermaphroditic or monoecious species. Self-fertilization rates varied inversely with population density and showed phylogenetic and geographic trends. The few direct measures of pollen flow (N=11 studies) suggest that pollen dispersal is widespread among low-density tropical trees, ranging from a mean of 200 m to over 19 km for species pollinated by small insects or bats. Future research needs to examine (1) the effect of inbreeding depression on observed outcrossing rates, (2) pollen dispersal in a wide range of pollination syndromes and ecological classes, (3) and the range of variation of mating system expression at different hierarchical levels, including individual, seasonal, population, ecological, landscape and range wide.

Genetic Markers↗

Mating systems of diploid and allotetraploid populations of tragopogon (Asteraceae). I. Natural populations

Although polyploidy is a significant force in the diversification of plants, the evolutionary consequences of polyploidization are not thoroughly understood. One possible consequence of polyploidy predicted by most population genetic theories is that the newly synthesized polyploid will self-fertilize at a greater rate than its diploid progenitors. To test for increased selfing rates in a polyploid, the mating systems of the allotetraploid Tragopogon mirus and one of its diploid progenitors, T. dubius, were compared. Tragopogon mirus is a recently derived species that arose sometime in the last 80 years and thus provides an opportunity to probe how quickly a shift in outcrossing rates might occur. Based on analyses of variation in maternal plants and their progeny arrays, the two tetraploid populations surveyed have higher outcrossing rates than the two diploid populations. This result is the opposite of that predicted by population genetic theory. This discrepancy between theoretical and empirical results may result from bias in the genetic sample, traits in the natural histories of the taxa involved or a lack of sufficient time since the formation of the polyploid (80 years or 40-80 generations) for a shift towards increased selfing to have occurred. Alternatively, the partial dominance model of inbreeding depression typically applied to polyploids may not be appropriate; the overdominance model predicts outcrossing rates in diploids and their tetraploid derivatives that are consistent with those observed in T. dubius and T. mirus.

Journal Article↗

Effect of population size on the mating system in a self-compatible, autogamous plant, aquilegia canadensis (Ranunculaceae)

In self-compatible plants, small populations may experience reduced outcrossing owing to decreased pollinator visitation and mate availability. We examined the relation between outcrossing and population size in eastern Ontario populations of Aquilegia canadensis. Experimental pollinations showed that the species is highly self-compatible, and can achieve full seed-set in the absence of pollinators via automatic self-pollination. We estimated levels of outcrossing (t) and parental inbreeding coefficients (F) from allozyme variation in naturally pollinated seed families for 10 populations ranging in size from 32 to 750 reproductive individuals. The proportion of seeds produced through outcrossing was generally low (mean = 0.29 +/- 0.02 SE) and varied widely among populations (range = 0.00-0.83). Accordingly, estimates of F were large (mean = 0.26 +/- 0.05) and significantly greater than zero in seven populations. As expected, four small populations (N < 40) outcrossed less (0.17 +/- 0.03) than six large populations (N > 90; 0.38 +/- 0. 03). However, parental plants were not significantly more inbred in small than large populations (P = 0.18). There was no difference in the germination of seeds from hand self- and cross-pollinations. However, population genetic estimates of inbreeding depression for survival expressed from seed to reproductive maturity were very high (mean delta = 1 - relative fitness of selfed seed = 0.88 +/- 0.14). The combination of self-compatibility and automatic self-pollination makes the mating system of A. canadensis sensitive to variation in ecological factors that affect the likelihood of cross-pollination.

Journal Article↗

Diallel crosses reveal patterns of variation in fruit-set, seed mass, and seed number in Asclepias incarnata.

Plants from each of two populations of Asclepias incarnata ssp. incarnata were hand-pollinated in diallel crosses and seed number per fruit, mean individual seed mass per fruit and fruit-set were determined for the parental plants. Two diallels were performed on the glasshouse-grown plants, one involving six plants from one population and the other involving five plants from the second population. The contributions to total phenotypic variation from joint, maternal, paternal, and two types of interaction sources of variation were then estimated. Variance attributable to maternal effects, reflecting differences between maternal plants resulting from environmental or genetic factors or both, comprised 31.3-68.4% of total variance in seed mass, 20.5-37.1% of variance in seed number, and 0.7-7.5% of variance in fruit-set. Interaction effects that depended on the direction of the cross accounted for 7.3-46.5% of variance in seed mass and 31.4-39.1% of variance in seed number. These interaction effects are consistent with the hypothesis that maternal plants provision resources differently to seeds depending on the specific genotypes of the embryos they contain. In the analysis of fruit-set, interaction effects independent of the direction of the cross (combined variance) explained 22.9-78.0% of phenotypic variance. These effects could have several causes, including inbreeding depression, shared incompatibility alleles, or 'control' of fruit-set by zygotes via hormonal signals or patterns of resource acquisition.

Phenotype↗