The effects of inheritance in tetraploids on genetic diversity and population divergence.

Polyploids are traditionally classified into allopolyploids and autopolyploids, based on their evolutionary origin and their disomic or multisomic mode of inheritance. Over the past decade it has become increasingly clear that there is a continuum between disomic and multisomic inheritance, with the rate of tetrasomy differing among species and among chromosomes within species. Here, we use a simple population genetic model to study the impact of the mode of inheritance on the genetic diversity and population divergence of tetraploids. We found that under almost strict disomic inheritance the tetraploid genome is divided into two separate subgenomes, such as found in classical allopolyploids. In those cases, assuming full tetrasomy in the analysis of polyploid genetic data will lead to an important bias in estimates of genetic diversity and population divergence. However, we found that even a low rate of allele exchange between the two subgenomes, at about one event per generation, is sufficient to homogenise the allele frequencies over the subgenomes, and the estimates become essentially unbiased. The inbreeding coefficient F(IS) can then be used to detect whether the estimates of diversity and divergence will be biased when full multisomy is assumed. Finally, we found that different summary statistics for measuring the strength of population differentiation are differentially affected by a deviation from full tetrasomy. Our model results provide several useful guidelines for the analysis of polyploid data, helping researchers to determine when their inferences are biased and which summary statistics to use.

Inheritance in tetraploid yeast revisited: segregation patterns and statistical power under different inheritance models.

In their recent article, Albertin et al. (2009) suggest an autotetraploid origin of 10 tetraploid strains of baker's yeast (Saccharomyces cerevisiae), supported by the frequent observation of double reduction meiospores. However, the presented inheritance results were puzzling and seemed to contradict the authors' interpretation that segregation ratios support a tetrasomic model of inheritance. Here, we provide an overview of the expected segregation ratios at the tetrad and meiospore level given scenarios of strict disomic and tetrasomic inheritance, for cases with and without recombination between locus and centromere. We also use a power analysis to derive adequate sample sizes to distinguish alternative models. Closer inspection of the Albertin et al. data reveals that strict disomy can be rejected in most cases. However, disomic inheritance with strong but imperfect preferential pairing could not be excluded with the sample sizes used. The possibility of tetrad analysis in tetraploid yeast offers a valuable opportunity to improve our understanding of meiosis and inheritance of tetraploids.

Male sterility in triploid dandelions: asexual females vs. asexual hermaphrodites.

Male reproductive output, pollen in plants and sperm in animals has been shown to constitute a substantial cost for many organisms. In parthenogenetic hermaphrodites, selection is therefore expected to reduce the allocation of resources to male reproductive output. However, sustained production of pollen or sperm has been observed in numerous asexual hermaphrodites. We studied the widespread production of pollen by triploid asexual dandelions, Taraxacum sect. Ruderalia, comparing rare male sterile individuals with pollen producing asexuals. We found that individuals can show plasticity in the production of pollen, but that it is nevertheless possible to distinguish between (facultatively) male sterile asexuals and male fertile asexuals. Based on evidence from genetic markers and crosses, we conclude that the male sterility in asexual dandelions is caused by nuclear genes, in contrast to the cytoplasmically inherited male sterility previously found in sexual dandelions. Male sterile lineages did not produce more seeds per flower head, heavier seeds or seeds that were more viable. However, male sterile plants did produce more seed heads and hence more seeds than pollen producing ones, indicating that they were able to reallocate resources toward seed production. Considering the difference in seed production, it remains puzzling that not more asexual dandelions are male sterile.

Nuclear-cytoplasmic male-sterility in diploid dandelions.

Male-sterility was found in diploid dandelions from two widely separated populations from France, and its inheritance was analysed by crossing a diploid male-sterile dandelion to diploid sexuals and triploid apomicts. Nuclear genetic variation, found in full-sib families, segregated for male-fertility, partial male-sterility, and full male-sterility, and also segregated for small-sized versus normally sized pollen. The crossing results are best explained by a cytoplasmic male-sterility factor in combination with two dominant restorer genes. Involvement of the cytoplasmic male-sterility factor was further investigated by chloroplast haplotyping. Male-sterility was exclusively associated with a rare chloroplast haplotype (designated 16b). This haplotype was found in seven male-sterile plants and one (apparently restored) male-fertile individual but does not occur in 110 co-existing male-fertile plants and not in several hundreds of individuals previously haplotyped. Apomicts with cytoplasmic male sterility were generated in some test crosses. This raises the question as to whether the male sterility found in natural dandelion apomicts, is of cytoplasmic or of nuclear genetic nature. As many breeding systems in Taraxacum are involved in shaping population structure, it will be difficult to predict the evolutionary consequences of nuclear-cytoplasmic male-sterility for this species complex.

Quantitative trait loci affecting growth-related traits in wild barley (Hordeum spontaneum) grown under different levels of nutrient supply.

The genetic basis of phenotypic plasticity of relative growth rate (RGR), its components and associated morphological traits was studied in relation to nutrient limitation. In all, 140 F(3) lines from a cross, made between two Hordeum spontaneum (wild barley) accessions sampled in Israel, were subjected to growth analysis under two nutrient levels. Quantitative trait loci (QTLs) were detected for RGR and three of its components, leaf area ratio (LAR), specific leaf area and leaf mass fraction (LMF). Indications for close linkage (potential pleiotropy) were found, for example, for LAR and LMF. An interesting case is on chromosome 6, at which QTLs for RGR and seed mass were detected in the same region. These QTLs had opposite additive effects, supporting earlier results that plants growing from lighter seeds had a higher RGR. Only two QTLs were significant under both nutrient conditions, suggesting large QTL x environment interactions for most traits. For 21 out of 26 QTLs, however, the additive genetic effect was of identical sign in both nutrient environments, but reached the significance threshold in only one of them. Nevertheless, some QTLs detected in one of the two environments had virtually no effect in the other, and QTLs for plasticity were detected for RGR, LAR and LMF, as well as for some morphological traits. QTLs with opposite effects under high and low nutrients were not found. Thus, at the genetic level, there was no evidence for a trade-off between faster growth at high versus low nutrient levels.

The relationship between relative growth rate and susceptibility to aphids in wild barley under different nutrient levels.

The Resource Availability Hypothesis (RAH) states that plants with a low Relative Growth Rate (RGR) and high levels of defence against herbivores or pathogens are favoured in habitats with low resource availability, whereas plants with a high potential RGR and low levels of defence are favoured in environments with high resource availability. High levels of defence are expected to result in lower reproduction and/or growth of the herbivores or pathogens. To test this hypothesis, four accessions of each of nine natural Hordeum spontaneum (wild barley) populations were grown in a climate chamber under two levels of nutrient supply. Susceptibility to Schizaphis graminum (greenbug) was quantified by placing a single adult greenbug on each plant and measuring its realised fecundity after 8 days. Data on potential RGR were available from a previous experiment. No support for the RAH was found. The correlation between potential RGR and greenbug reproduction was not significant, neither at the high nor at the low level of nutrient supply. Furthermore, on average plants grown under high and low nutrients did not differ in susceptibility. However, accessions-within-populations differed in the way susceptibility was affected by nutrient supply, and most accessions had a higher susceptibility under nutrient-poor conditions. It could be that these accessions differed in the spectrum of secondary metabolites they produced. Whatever the cause, the genetic variation for the reaction in susceptibility to nutrient supply suggests that selection could act in favour of more or less plasticity in plants without any apparent change in potential RGR.

Adaptive phenotypic plasticity: consensus and controversy.

Phenotypic plasticity is an environmentally based change in the phenotype. Understanding the evolution of adaptive phenotypic plasticity has been hampered by dissenting opinions on the merits of different methods of description, on the underlying genetic mechanisms, and on the way that plasticity is affected by natural selection in a heterogeneous environment. During much of this debate, the authors of this article have held opposing views. Here, we attempt to lay out current issues and summarize the areas of consensus and controversy surrounding the evolution of plasticity and the reaction norm (the set of phenotypes produced by a genotype over a range of environments).

Selection on reaction norms, genetic correlations and constraints.

Two approaches to the evolution of phenotypic plasticity in heterogeneous environments have recently been put forward. The first focuses on selection on the character expression within each environment; plasticity is seen as a by-product of local selection in various habitats. The second approach focuses on selection on the parameters of the response function of genotypes, and selection is thought to change the frequencies of 'plasticity' genes that affect the function. This paper discusses the relationship between the two approaches, with emphasis on applications. A method is described that allows switching from one approach to the other. It is argued that character state and reaction norm approaches, while to a large extent interchangeable, usually differ in the response function chosen. This choice, however, may strongly affect the biological interpretation. The methods outlined in this paper permit one to look at the data from different perspectives in order to avoid this danger.

Ontoecogenophyloconstraints? The chaos of constraint terminology.

The world of evolutionary biology today is being bombarded with all kinds of possible constraints to the process of natural selection. Are we witnessing the end of the neodarwinistic theory of evolution, as some may like to see it, or is it just another whim of giving new names to old things? Here, we attempt to unravel the meaning and name-giving of constraints in a small and nonrandom sample of the literature, and suggest a way out from the present confusion of usages.

Within-population variability in morphology and life history of Plantago major L. ssp. pleiosperma Pilger in relation to environmental heterogeneity.

An attempt was made to relate variation in life-history characteristics within a population of Plantago major ssp. pleiosperma to small-scale environmental variability. At a beach plain, embanked in 1966, a mosaic environment was distinguished with spatial variability in vegetation structure as well as in nutrient availability and water content of the soil. Differences between three subsites in comtemporary selection were demonstrated, e.g. in shoot morphology and allocation to reproductive tissue. The effects of nutrient supply and waterlogging on morphology and life history were studied on lines from the three subsites in a greenhouse. For most of the traits high levels of phenotypic plasticity were observed, covering almost entirely the observed phenotypic variability at the beach plain. In all treatments lines from the shrubs had, however, a higher leaf-area ratio as well as delayed flowering when compared to lines from more open subsites. In addition, in a reciprocal transplant experiment it was demonstrated that lines from the shrubs had larger shoots with e.g. broader leaves in the shady environment of the shrubs than other lines.From the experiments no indications were obtained that lines from any subsite were especially adapted to specific levels of nutrient supply or water content of the soil. With respect to these environmental factors P. major ssp. pleisoperma might occur and reproduce at all subsites by means of phenotypic plasticity, e.g. in plant form. However, it is suggested that spatial variability in vegetation structure caused a population subdivision in allocation patterns, leaf form and life history at the beach plain, over distances of about 15-25 m. This differentation occurred during primary succession over a period of twenty years.