Performance of aneuploid backcross hybrids between the crop Brassica napus and its wild relative B. rapa.

Crossings between the diploid wild Brassica rapa (AA, 2n = 20) and the tetraploid cultivar B. napus (AACC, 2n = 38) can readily be made. Backcrosses to the wild B. rapa (BC1 ) produce aneuploids with variable chromosome numbers between 20 and 29. How does survival and performance relate to DNA content of plants? Growth of the BC1 plants was measured in the lab. One plant in the F1 self-pollinated spontaneously and produced abundant F2 seeds that were also examined. The number of C-chromosomes was estimated from DNA values obtained with flow cytometry. Average DNA value of the BC1 was similar to that of the parents, which shows that C-chromosomes do not reduce success of pollen or embryos. The average DNA value in the F2 was 13% higher than in the F1 , suggesting that extra C-chromosomes facilitated gamete success and/or embryo survival. Under both optimal and drought stress conditions growth and survival of BC1 hybrids was similar to that of B. rapa. No significant correlations existed between growth or survival and DNA value. Aneuploid plants were not inferior under the conditions of the growth room and may persist in nature. We discuss other factors, such as herbivory, that could prevent hybrid establishment in the field.

Hybridisation and introgression between Brassica napus and B. rapa in the Netherlands.

We used flow cytometry, chromosome counting and AFLP markers to investigate gene flow from the crop plant oilseed rape, Brassica napus (AACC) to wild B. rapa (AA) in the Netherlands. From 89 B. napus source populations investigated, all near cropping fields or at transhipment sites, only 19 contained a B. rapa population within a 2.5-km radius. During our survey we found only three populations with F1 hybrids (AAC), as recognized by their nine extra chromosomes and by flow cytometry. These hybrids were all collected in mixed populations where the two species grew in close proximity. Populations with F1 hybrids were not close to crops, but instead were located on road verges with highly disturbed soils, in which both species were probably recruited from the soil seed bank. Many plants in the F2, BC1 or higher backcrosses are expected to carry one to eight C chromosomes. However, these plants were not observed among the hybrids. We further investigated introgression with molecular markers (AFLP) and compared sympatric B. rapa populations (near populations of B. napus) with control populations of B. rapa (no B. napus within at least 7 km). We found no difference between sympatric and control populations in the number of C markers in B. rapa, nor did we find that these sympatric populations closely resembled B. napus. Our data show that hybrids occur but also suggest no recent introgression of alleles from the crop plant B. napus into wild B. rapa in the Dutch populations studied.

Morphological markers for the detection of introgression from cultivated into wild carrot (Daucus carota L.) reveal dominant domestication traits.

Hybridisation and subsequent introgression have recently received much attention in the context of genetically modified crops. But crop-wild hybrid detection in the field can be difficult, as most domestication traits seem to be recessive, and the hybrid phenotype may also depend on the direction of the cross or environmental factors. Our aim was to develop a reliable set of morphological markers that differ between two wild and 13 cultivated carrots (Daucus carota L.) and to evaluate their inheritance in hybrid lines. We then examined these morphological markers in four F1 hybrids obtained by fertilising plants from the two wild accessions with pollen from two common carrot cultivars. Of the 16 traits that differed between the two carrot subspecies, three took intermediate values in the hybrids, eight resembled the cultivar parent (dominant domestication traits), two resembled the wild parent (domestication traits recessive), and three were not significant or growth condition-dependent. Root:shoot ratio was seven times higher for cultivars than for wild plants, while still attaining equivalent total dry weight, which shows that dry matter production by the shoot is much higher in cultivars than in wild plants. High root:shoot ratios were also present in the hybrids. While we found no maternal effects, the type of cultivar used for pollination had an impact on hybrid characteristics. The morphological markers developed here provide insights into the mode of inheritance of ecologically relevant traits and can be useful for pre-screening wild populations for hybrid detection prior to genetic analysis.

Metabolic fingerprinting reveals differences between shoots of wild and cultivated carrot (Daucus carota L.) and suggests maternal inheritance or wild trait dominance in hybrids.

Differences between the metabolic content of cultivars and their related wild species not only have implications for breeding and food quality, but also for the increasingly studied area of crop to wild introgression. Wild and cultivated western carrots belong to the same outcrossing species and hybridize under natural conditions. The metabolic fingerprinting of Dutch wild carrot and of western orange carrot cultivar shoots using (1)H NMR showed only quantitative differences in chemical content, indicating relatively low divergence after domestication. Main differences reside in the primary metabolite content and in the concentrations of chlorogenic acid and feruloyl quinic acid in the shoots of the different carrot types. Wild×cultivar hybrids cannot be distinguished from wild plants based on the metabolome, suggesting maternal, maternal environment, or dominance effects, and indicating high hybrid fitness in wild conditions. Considering these similarities, introgression is a real possibility in carrots, but understanding its consequences would require further studies using backcrosses in a multiple environments.

Paternal effects on seed mass in Arabidopsis thaliana.

Who is in control of seed size, and do some fathers sire bigger seeds than others? We used isogenic male-sterile genotypes of the Arabidopsis thaliana accessions Col and Ler. By fertilising flowers side-by-side with either pollen from the same accession ('self-pollination') or pollen from another accession (outcrossing), we compared, on the same mother plant, seed set of flowers that were very similar in resource status. Some paternal genotypes had a significant effect on seed mass, with the most extreme father siring seeds 15.3% heavier than seeds resulting from 'self-pollination'. There was no correlation between seed mass of paternal parents and the seeds they sired. We discuss the evolution of seed size as a tug-of-war between parent and offspring.

Inheritance of progeny sex ratio in Urtica dioica.

Seed samples collected from female Urtica dioica plants in the field showed considerable inter-family variation in the sex ratio (faction of males). To investigate the inheritance pattern of the sex ratio trait, crosses were performed between individual male and female plants from different sex ratio families. Our results suggest, at least for the families studied here, that maternal parents strongly contribute to the variation in the primary sex ratio. Furthermore, progeny sex ratios from reciprocal crosses were significantly different and resembled the sex ratios produced by their maternal parents. We discuss the possible mechanisms underlying maternal control.

Herbivory and abiotic factors affect population dynamics of Arabidopsis thaliana in a sand dune area.

Population dynamics of the annual plant Arabidopsis thaliana (L.) Heynh. were studied in a natural habitat of this species on the coastal dunes of the Netherlands. The main objective was to elucidate factors controlling population dynamics and the relative importance of factors affecting final population density. Permanent plots were established and plants were mapped to obtain data on survival and reproductive performance of each individual, with special attention to herbivore damage. In experimental plots we studied how watering, addition of nutrients, artificial disturbance, and natural herbivores affected survival and growth. Mortality was low during autumn and early winter and high at the time of stem elongation, between February and April. A key factor analysis showed a high correlation between mortality from February to April and total mortality. The specialist weevils Ceutorhyncus atomus and C. contractus (Curculionidae) were identified as the major insect herbivores on A. thaliana, reducing seed production by more than 40 %. These herbivores acted in a plant size-dependent manner, attacking a greater fraction of the fruits on large plants. While mortality rates were not affected by density, fecundity decreased with density, although the effect was small. Adding water reduced mortality in rosette and flowering plant stages. Soil disturbance did not increase seed germination, but did have a significant positive effect on survival of rosette and flowering plants. Seed production of A. thaliana populations varied greatly between years, leading to population fluctuations, with a small role for density-dependent fecundity and plant size-dependent herbivory.

Heritable variation in seed sex ratio of the stinging nettle (Urtica dioica).

Male and female flowering plants of the dioecious Urtica dioica occur in approximately equal numbers in our study area on the coastal sand dunes of Meijendel. The seed sex ratio (SSR, fraction of males) collected from female plants in the field varied between 0.05 and 0.76, and differed significantly between maternal parents. After one generation of selection for either high or low SSR, female plants produced seed batches with sex ratios as extreme as 0.08 and 0.73. Natural populations of U. dioica harbour considerable genetic variation in SSR.

Hamilton's rule, imprinting and parent-offspring conflict over seed mass in partially selfing plants.

When genes in the offspring control the provisioning of the seed, the optimal seed size can be calculated exactly by applying Hamilton's rule. When seed size is a compromise between mother and offspring, we predict that outcrossing plant species produce larger seeds than selfers. This trend was found in the British flora and in a number of well-studied plant families. The analysis was extended to imprinting, a conditional strategy in which a gene in the offspring takes more resources when derived from the father than from the mother. The conditions for imprinting to be selected were rather restrictive. The analysis is relevant for the current debate about the evolution of imprinting in Arabidopsis thaliana.

Sex ratio of some long-lived dioecious plants in a sand dune area.

In dioecious plants the fraction of males among flowering plants in the field (the secondary sex ratio) is the result of the fraction of males in the seeds (the primary sex ratio) and the subsequent survival and age at first reproduction of the two genders. It has been assumed that survival and age at first reproduction are the main determinants of biased secondary sex ratio but, especially for long-lived perennials, few data are available. We address this issue for natural populations of four long-lived perennials in a dune area. In Asparagus officinale and Bryonia dioica, the secondary sex ratio was unbiased. In Salix repens the secondary sex ratio was female-biased (0.337). Hippophae rhamnoides populations were male-biased; the average sex ratio of flowering plants was 0.658, while the fraction of males varied between 0.39 near the sea to 0.84 at the inland side of the dunes. The primary sex ratio was estimated by germinating seeds and growing plants under favourable conditions with minimal mortality. In S. repens the primary sex ratio in seeds was variable among mother plants and was, on average, female-biased (0.289). This is close to the secondary sex ratio, suggesting that the female bias already originates in the seed stage. In Hippophae rhamnoides the primary sex ratio was slightly male-biased (0.564). We argue that in this species, apart from the primary sex ratio, higher mortality and a later age at first reproduction for females contribute to the strong male bias among flowering plants in the field.

Why girls want to be boys.

The mechanisms by which sex is genetically determined are bewilderingly diverse and appear to change rapidly during evolution.(1) What makes the sex-determining process so prone to perturbations? Two recent articles(2,3) explore theoretically the role of genetic conflict in sex determination evolution. Both studies use the idea that selection on sex-determining genes may act differently in parents and in offspring and they suggest that the resulting conflict can drive changes in sex-determining mechanisms.

Direct and indirect estimates of the selfing rate in small and large individuals of the bumblebee pollinated Cynoglossum officinale L (Boraginaceae).

We measured the relationship between selfing rates and flower number in an experimental population of bumblebee pollinated Cynoglossum officinale, with plants differing in flower number. Results were compared with the prediction of a model based on pollen dynamics and pollinator behaviour. The selfing rate, as measured by multilocus oligonucleotide DNA fingerprinting, increased with flower number and ranged from 0% to 70%. Flowers on large plants received an equal number of visits from bumblebees as flowers on small plants. On large plants more flowers in a row were visited, inducing geitonogamy. The overall relationship between selfing rate and number of flowers can be explained by pollen dynamics and pollinator behaviour without invoking postpollination processes such as differential pollen tube growth and abortion.

Sex and size in cosexual plants.

There are conceptual and practical difficulties in measuring the exact shape of fitness-gain curves and sex allocation, and these hamper empirical testing of some of the basic predictions of sex allocation theory for plants. Nevertheless, our knowledge of the processes that shape fitness-gain curves allows us to formulate hypotheses to test predictions of sex allocation theory. One such hypothesis is that plants adjust their gender according to size. The connection between plant size and gender was generally thought to be weak. Recent data, however, suggest that size-dependent sex allocation (SDS) is a common phenomenon in hermaphrodites and other cosexual plants.

Geitonogamy: The neglected side of selfing.

Flowers of many angiosperm species are well adapted to avoid self-pollination, for instance by temporal and spatial separation of pollen and stigma within the same flower. However, such adaptations do not prevent the transfer of pollen between flowers on the same plant (geitonogamy). Recent empirical studies, aided by advances in field techniques, statistical methods and modelling, show that geitonogamy often is substantial and increases with plant size. Selfing by geitonogamy incurs a fitness cost of reduced pollen export, and recent reports show that seed set can suffer as well, even in self-incompatible species. Geitonogamy has important implications for sex-allocation theory, the evolution of dioecy and other issues in evolutionary biology.

The efficacy of dispersal in relation to safe site area and seed production.

A model is developed to define the efficacy of dispersal in relation to safe site area and seed production. Efficacy is measured as the expected number of progeny from one parent. It is shown that maximization of efficacy does not depend on the density of safe sites. When safe sites are confined to a restricted area around the parent, and safe sites are small or few propagules are produced, dispersal curves with short tails are most efficacious; when safe sites are larger or when more propagules are produced, distributions with longer tails become more advantageous.

What are the advantages of dispersing; a paper by Kuno explained and extended.

Contrary to Kuno's (1981) contention, dispersing does not help and individual to get a larger average progeny in an unpredictable and heterogeneous but nonlimiting environment: average progeny is exactly equal for (partially) dispersing and nondispersing populations. However, the geometric time averages of pro-capita reproduction as well as geometric averages over replicates of final progeny size after a fixed number of years differ, just as Kuno asserts. Moreover, if populations of the two types are grown in mixed culture it is the disperser who will win in the long run. This even applies if dispersal means the incurring of some additional mortality. Models with partial dispersal are much more complicated to deal with than models with either a complete redistribution each generation or no dispersal at all, contrary to the assertion of e.g. Venable and Lawlor (1980). Partial dispersers will win from nondispersers, but the optimal amount of dispersal unfortunately seems to depend sensitively on the details of the model specification, except that it has to be small if the number of independent patches is large.