The homomorphic self-incompatibility system in Oleaceae is controlled by a hemizygous genomic region expressing a gibberellin pathway gene.

In flowering plants, outcrossing is commonly ensured by self-incompatibility (SI) systems. These can be homomorphic (typically with many different allelic specificities) or can accompany flower heteromorphism (mostly with just two specificities and corresponding floral types). The SI system of the Oleaceae family is unusual, with the long-term maintenance of only two specificities but often without flower morphology differences. To elucidate the genomic architecture and molecular basis of this SI system, we obtained chromosome-scale genome assemblies of Phillyrea angustifolia individuals and related them to a genetic map. The S-locus region proved to have a segregating 543-kb indel unique to one specificity, suggesting a hemizygous region, as observed in all distylous systems so far studied at the genomic level. Only one of the predicted genes in this indel region is found in the olive tree, Olea europaea, genome, also within a segregating indel. We describe complete association between the presence/absence of this gene and the SI types determined for individuals of seven distantly related Oleaceae species. This gene is predicted to be involved in catabolism of the gibberellic acid (GA) hormone, and experimental manipulation of GA levels in developing buds modified the male and female SI responses of the two specificities in different ways. Our results provide a unique example of a homomorphic SI system, where a single conserved gibberellin-related gene in a hemizygous indel underlies the long-term maintenance of two groups of reproductive compatibility.

Widespread coexistence of self-compatible and self-incompatible phenotypes in a diallelic self-incompatibility system in Ligustrum vulgare (Oleaceae).

The breakdown of self-incompatibility (SI) in angiosperms is one of the most commonly observed evolutionary transitions. While multiple examples of SI breakdown have been documented in natural populations, there is strikingly little evidence of stable within-population polymorphism with both inbreeding (self-compatible) and outcrossing (self-incompatible) individuals. This absence of breeding system polymorphism corroborates theoretical expectations that predict that in/outbreeding polymorphism is possible only under very restricted conditions. However, theory also predicts that a diallelic sporophytic SI system should facilitate the maintenance of such polymorphism. We tested this prediction by studying the breeding system of Ligustrum vulgare L., an insect-pollinated hermaphroditic species of the Oleaceae family. Using stigma tests with controlled pollination and paternity assignment of open-pollinated progenies, we confirmed the existence of two self-incompatibility groups in this species. We also demonstrated the occurrence of self-compatible individuals in different populations of Western Europe arising from a mutation affecting the functioning of the pollen component of SI. Our results show that the observed low frequency of self-compatible individuals in natural populations is compatible with theoretical predictions only if inbreeding depression is very high.

Diallelic self-incompatibility is the main determinant of fertilization patterns in olive orchards.

Self-incompatibility (SI) in flowering plants potentially represents a major obstacle for sexual reproduction, especially when the number of S-alleles is low. The situation is extreme in the commercially important olive tree, where in vitro pollination assays suggested the existence of a diallelic SI (DSI) system involving only two groups (G1 and G2). Varieties belonging to the same SI group cannot fertilize each other, such that successful fruit production is predicted to require pollination between varieties of different groups. To test this prediction, we explored the extent to which the DSI system determines fertilization patterns under field conditions. One hundred and seventeen olive cultivars were first genotyped using 10 highly polymorphic dinucleotide Simple Sequence Repeat (SSR) markers to ascertain varietal identity. Cultivars were then phenotyped through controlled pollination tests to assign each of them to one of the two SI groups. We then collected and genotyped 1440 open pollinated embryos from five different orchards constituted of seven local cultivars with known group of incompatibility groups. Embryos genotype information were used: (i) to assign embryos to the most likely pollen donor genotype in the neighbourhood using paternity analysis, and (ii) to compare the composition of the pollen cloud genetic among recipient trees in the five sites. The paternity analysis showed that the DSI system is the main determinant of fertilization success under field open pollination conditions: G1 cultivars sired seeds exclusively on G2 cultivars, and reciprocally. No self-fertilization events were observed. Our results demonstrate that DSI is a potent force determining pollination success among varieties within olive orchards used for production. They have the potential to improve management practices by guiding the selection of compatible varieties to avoid planting orchards containing sets of varieties with strongly unbalanced SI groups, as these would lead to suboptimal olive production.

Effect of preferred walking speed on the upper body range of motion and mechanical work during gait before and after spinal fusion for patients with idiopathic scoliosis.

BACKGROUND: Scoliosis may have an effect on gait parameters, the kinematics of the lower limbs and the spine, and mechanical work with specific gait speed. Imposed gait speed may influence these effects. Following spinal fusion in the case of idiopathic scoliosis, patients fear subsequent and considerable back stiffness and kinetic consequences. The aim of this retrospective study was to evaluate the upper body range of motion and mechanical work before and after spinal fusion in of free gait speed conditions. METHODS: Twenty-two patients with idiopathic scoliosis and twenty-two asymptomatic controls were included. Patients were analyzed before and one year after spinal fusion. Based on full body modeling and motion capture, we measured gait speed, cadence, stride length, the mobility of the upper and lower spinal segments (in each plane), and mechanical work (with and without dimensionless scaling strategy). FINDINGS: Patients walked significantly slower than controls. The same speed was noticed before and after fusion. Only the lower back kinematics in the frontal plane was reduced before fusion. Spinal fusion further reduced the mobility of the pelvis segment in the sagittal plane in comparison to controls. Scaling external work was associated with higher values for patients. INTERPRETATION: Spinal fusion improves pelvic and thorax-pelvis mobility (during the stance phase) in the frontal plane. The impact of scoliosis on the upper body range of motion was limited on the thorax-pelvis, corresponding to a stiffening effect. With such restrictions, an increase in normalized external work was observed for similar normalized internal work.

Genetic Mapping of the Incompatibility Locus in Olive and Development of a Linked Sequence-Tagged Site Marker.

The genetic control of self-incompatibility (SI) has been recently disclosed in olive. Inter-varietal crossing confirmed the presence of only two incompatibility groups (G1 and G2), suggesting a simple Mendelian inheritance of the trait. A double digest restriction associated DNA (ddRAD) sequencing of a biparental population segregating for incompatibility groups has been performed and high-density linkage maps were constructed in order to map the SI locus and identify gene candidates and linked markers. The progeny consisted of a full-sib family of 229 individuals derived from the cross 'Leccino' (G1) × 'Dolce Agogia' (G2) varieties, segregating 1:1 (G1:G2), in accordance with a diallelic self-incompatibility (DSI) model. A total of 16,743 single nucleotide polymorphisms was identified, 7,006 in the female parent 'Leccino' and 9,737 in the male parent 'Dolce Agogia.' Each parental map consisted of 23 linkage groups and showed an unusual large size (5,680 cM in 'Leccino' and 3,538 cM in 'Dolce Agogia'). Recombination was decreased across all linkage groups in pollen mother cells of 'Dolce Agogia,' the parent with higher heterozygosity, compared to megaspore mother cells of 'Leccino,' in a context of a species that showed exceptionally high recombination rates. A subset of 109 adult plants was assigned to either incompatibility group by a stigma test and the diallelic self-incompatibility (DSI) locus was mapped to an interval of 5.4 cM on linkage group 18. This region spanned a size of approximately 300 Kb in the olive genome assembly. We developed a sequence-tagged site marker in the DSI locus and identified five haplotypes in 57 cultivars with known incompatibility group assignment. A combination of two single-nucleotide polymorphisms (SNPs) was sufficient to predict G1 or G2 phenotypes in olive cultivars, enabling early marker-assisted selection of compatible genotypes and allowing for a rapid screening of inter-compatibility among cultivars in order to guarantee effective fertilization and increase olive production. The construction of high-density linkage maps has led to the development of the first functional marker in olive and provided positional candidate genes in the SI locus.

Polygamy or subdioecy? The impact of diallelic self-incompatibility on the sexual system in Fraxinus excelsior (Oleaceae).

How flowering plants have recurrently evolved from hermaphroditism to separate sexes (dioecy) is a central question in evolutionary biology. Here, we investigate whether diallelic self-incompatibility (DSI) is associated with sexual specialization in the polygamous common ash (Fraxinus excelsior), which would ultimately facilitate the evolution towards dioecy. Using interspecific crosses, we provide evidence of strong relationships between the DSI system and sexual phenotype. The reproductive system in F. excelsior that was previously viewed as polygamy (co-occurrence of unisexuals and hermaphrodites with varying degrees of allocation to the male and female functions) and thus appears to actually behave as a subdioecious system. Hermaphrodites and females belong to one SI group and functionally reproduce as females, whereas males and male-biased hermaphrodites belong to the other SI group and are functionally males. Our results offer an alternative mechanism for the evolution of sexual specialization in flowering plants.

Controlling for genetic identity of varieties, pollen contamination and stigma receptivity is essential to characterize the self-incompatibility system of Olea europaea L.

Bervillé et al. express concern about the existence of the diallelic self-incompatibility (DSI) system in Olea europaea, mainly because our model does not account for results from previous studies from their group that claimed to have documented asymmetry of the incompatibility response in reciprocal crosses. In this answer to their comment, we present original results based on reciprocal stigma tests that contradict conclusions from these studies. We show that, in our hands, not a single case of asymmetry was confirmed, endorsing that symmetry of incompatibility reactions seems to be the rule in Olive. We discuss three important aspects that were not taken into account in the studies cited in their comments and that can explain the discrepancy: (i) the vast uncertainty around the actual genetic identity of vernacular varieties, (ii) the risk of massive contamination associated with the pollination protocols that they used and (iii) the importance of checking for stigma receptivity in controlled crosses. These studies were thus poorly genetically controlled, and we stand by our original conclusion that Olive tree exhibits DSI.

Elucidation of the genetic architecture of self-incompatibility in olive: Evolutionary consequences and perspectives for orchard management.

The olive (Olea europaea L.) is a typical important perennial crop species for which the genetic determination and even functionality of self-incompatibility (SI) are still largely unresolved. It is still not known whether SI is under gametophytic or sporophytic genetic control, yet fruit production in orchards depends critically on successful ovule fertilization. We studied the genetic determination of SI in olive in light of recent discoveries in other genera of the Oleaceae family. Using intra- and interspecific stigma tests on 89 genotypes representative of species-wide olive diversity and the compatibility/incompatibility reactions of progeny plants from controlled crosses, we confirmed that O. europaea shares the same homomorphic diallelic self-incompatibility (DSI) system as the one recently identified in Phillyrea angustifolia and Fraxinus ornus. SI is sporophytic in olive. The incompatibility response differs between the two SI groups in terms of how far pollen tubes grow before growth is arrested within stigma tissues. As a consequence of this DSI system, the chance of cross-incompatibility between pairs of varieties in an orchard is high (50%) and fruit production may be limited by the availability of compatible pollen. The discovery of the DSI system in O. europaea will undoubtedly offer opportunities to optimize fruit production.

Evidence for the long-term maintenance of a rare self-incompatibility system in Oleaceae.

A rare homomorphic diallelic self-incompatibility (DSI) system discovered in Phillyrea angustifolia (family Oleaceae, subtribe Oleinae) can promote the transition from hermaphroditism to androdioecy. If widespread and stable in Oleaceae, DSI may explain the exceptionally high rate of androdioecious species reported in this plant family. Here, we set out to determine whether DSI occurs in another Oleaceae lineage. We tested for DSI in subtribe Fraxininae, a lineage that diverged from subtribe Oleinae c. 40 million yr ago. We explored the compatibility relationships in Fraxinus ornus using 81 hermaphrodites and 25 males from one natural stand and two naturalized populations using intra- and interspecific stigma tests performed on F. ornus and P. angustifolia testers. We uncovered a DSI system with hermaphrodites belonging to one of two self-incompatibility (SI) groups and males compatible with both groups, making for a truly androdioecious reproductive system. The two human-founded populations contained only one of the two SI groups. Our results provide evidence for the evolutionary persistence of DSI. We discuss how its stability over time may have affected transitions to other sexual systems, such as dioecy.

Selfish male-determining element favors the transition from hermaphroditism to androdioecy.

According to the current, widely accepted paradigm, the evolutionary transition from hermaphroditism toward separate sexes occurs in two successive steps: an initial, intermediate step in which unisexual individuals, male or female, sterility mutants coexist with hermaphrodites and a final step that definitively establishes dioecy. Two nonexclusive processes can drive this transition: inbreeding avoidance and reallocation of resources from one sexual function to the other. Here, we report results of controlled crosses between males and hermaphrodites in Phillyrea angustifolia, an androdioecious species with two mutually intercompatible, but intraincompatible groups of hermaphrodites. We observed different segregation patterns that can be explained by: (1) epistatic interactions between two unlinked diallelic loci, determining sex and mating compatibility, and (2) a mutation with pleiotropic effects: female sterility, full compatibility of males with both hermaphrodite incompatibility groups, and complete male-biased sex-ratio distortion in one of the two groups. Modeling shows that these mechanisms can explain the high frequency of males in populations of P. angustifolia and can promote the maintenance of androdioecy without requiring inbreeding depression or resource reallocation. We thus argue that segregation distortion establishes the right conditions for the evolution of cryptic dioecy and potentially initiates the evolution toward separate sexes.

The joint evolution and maintenance of self-incompatibility with gynodioecy or androdioecy.

Mating systems show two kinds of frequent transitions: from hermaphroditism to dioecy, gynodioecy or androdioecy, or from self-incompatibility (SI) to self-compatibility (SC). While models have mostly investigated these two kinds of transitions as independent, empirical observations suggest that, to some extent, they can evolve jointly. Here, we study the joint evolution and maintenance of SI and androdioecy or SI and gynodioecy by the means of phenotypic models. Our models focus on three parameters: the unisexuals׳ advantage relative to that of the hermaphrodites due to resource reallocation, inbreeding depression and the selfing rate. We assume no pollen limitation or discounting. We show that SI helps the maintenance of androdioecy, but favors the loss of gynodioecy, and also that androdioecy facilitates the maintenance of SI, whereas gynodioecy does not affect it. We finally investigate how gynodioecy and androdioecy may affect the diversification of SI groups, especially considering an evolutionary pathway through SC intermediates. We show that while androdioecy prevents the increase of the number of SI groups, under certain conditions of inbreeding depression and selfing rates, gynodioecy allows it.

Evolution of a genetic polymorphism with climate change in a Mediterranean landscape.

Many species show changes in distribution and phenotypic trait variation in response to climatic warming. Evidence of genetically based trait responses to climate change is, however, less common. Here, we detected evolutionary variation in the landscape-scale distribution of a genetically based chemical polymorphism in Mediterranean wild thyme (Thymus vulgaris) in association with modified extreme winter freezing events. By comparing current data on morph distribution with that observed in the early 1970s, we detected a significant increase in the proportion of morphs that are sensitive to winter freezing. This increase in frequency was observed in 17 of the 24 populations in which, since the 1970s, annual extreme winter freezing temperatures have risen above the thresholds that cause mortality of freezing-sensitive morphs. Our results provide an original example of rapid ongoing evolutionary change associated with relaxed selection (less extreme freezing events) on a local landscape scale. In species whose distribution and genetic variability are shaped by strong selection gradients, there may be little time lag associated with their ecological and evolutionary response to long-term environmental change.

A self-incompatibility system explains high male frequencies in an androdioecious plant.

Androdioecy is a sexual system in which males co-occur with hermaphrodites, which have both male and female function. Stable androdioecy is rare in nature, and theory suggests that it requires that males sire more than twice as many offspring as hermaphrodites. In several members of the olive family (Oleaceae), androdioecy occurs with higher frequencies of males than predicted by theory. In Phillyrea angustifolia L., we found that high male frequencies can be maintained in natural populations because hermaphrodites belong exclusively to one of two self-incompatibility groups, and thus, each can fertilize only half of all pollen recipients. In contrast, males can pollinate all hermaphrodites. Thus, in this species, the reproductive disadvantage that males face due to the loss of female function is offset by the fact that all males are fully compatible with all pollen recipients.

[Screening and treatment of malnutrition: European Council Resolution and its potential application in Switzerland]. / Dépistage et traitement de la dénutrition: Résolution du Conseil de l'Europe et potentiel d'application en Suisse.

The deleterious consequences of hospital malnutrition are progressively being recognized by the public. The Committee of the Ministers of the European Council has released a resolution for food and nutrition and ask the state members to apply national recommendations in their hospitals. The evaluated domains and actions to be undertaken are based on 4 elements: Access to a healthy and varied food supply is a fundamental human right; a high number of patients hospitalized in Europe are malnourished (30-60%); the demonstration that malnutrition of hospitalized patients increases length of hospital stay and medical costs; the demonstrated beneficial effects of a hospital food service and of optimal nutritional care on morbidity, healing and quality of life. The resolution indicates clearly the options to be followed to optimize the actions and decisions at three levels: patient care, support services, and administration. All caregivers must be involved in the implementation of the recommendations in a way to adapt them to the local needs and restrictions. This review summarizes the actions to be taken by medical and other caregivers in order to sensitize them to the above-mentioned problems and to initiate practical options to treat malnourished patients or patients at risk of becoming malnourished.

THE EVOLVING GENETIC HISTORY OF A POPULATION OF LATHYRUS SYLVESTRIS: EVIDENCE FROM TEMPORAL AND SPATIAL GENETIC STRUCTURE.

We analyze patterns of genetic microdifferentiation within a natural population of Lathyrus sylvestris, a perennial herb with both sexual reproduction and clonal growth. In a population from the northern foothills of the Pyrénées in southwestern France, a combined demographic and genetic investigation enabled the study not only of spatial genetic structure of the population, but also of the history of the population's spatial genetic structure over time. Excavation of all individuals allowed identification of clonemates. Age of each individual was determined by counting annual growth rings in the taproot, a method tested with individuals of known age planted in experimental gardens. Each individual was mapped, and genotypes of all individuals were determined using allozyme markers for a number of polymorphic loci. Distribution patterns and spatial genetic structure, both for all individuals and for different age classes, were analyzed using spatial autocorrelation statistics (Geary's Index, Moran's Index). Patterns of gene flow within the population were also studied using F-statistics and tests for random associations of alleles. Because age, allele frequencies, and location were known for each individual, it was possible to study how spatial genetic structure changed over time. Results from all these diverse approaches are consistent with one another, and clearly show the following: (1) founder effects, with the study transect being first colonized by individuals at either end of the transect that were homozygous for different alleles at one marker locus; (2) a difference in spatial distribution of individuals originated from sexual reproduction (seedlings) and from clonal growth (connected individuals); (3) restricted gene flow, due to inbreeding among related, clumped individuals; and (4) increase in heterozygote deficit within the youngest cohort of individuals. The results indicate that genetic differentiation in time was much less marked than differentiation in space. Nevertheless, the results revealed that the studied population is experiencing demographic and genetic variation in time, suggesting that it is not at equilibrium. On the one hand, spatial structuring is becoming less marked due to the recombination of founder genotypes; on the other hand, as establishment of new individuals increases, a new spatial structure emerges due to mating between relatives.

Interference between Hieracium pilosella and Arrhenatherum elatius in colliery spoils of north of France : Allelopathy or competition?

On colliery heaps in northern France, a tall grassland community dominated by Arrhenatherum elatius, give place to a thin grassland community in which Hieracium pilosella is very abundant. It has been claimed that Hieracium pilosella is an allelopathic species and this phenomenon has been investigated as an explanation of this phase of plant succession. The importance of osmotic pressure in tested plant extracts is demonstrated and may be responsible for presumed allelopathy. Two phytotoxic compounds have been revealed in roots+rhizomes of Hieracium pilosella (umbelliferon and apigenin-glucoside). A third phytotoxic compound was exuded by roots in hydroponic cultures (7-glucoside-umbelliferon or skimin). However no toxicity of soil has been found and no phytotoxic compounds appeared to be present in the soil under Hieracium pilosella. Experimental mixed cultures (sterile or non-sterile conditions) reveal suppression of Hieracium pilosella by Arrhenatherum elatius rather than the reverse. Allelopathy cannot be invoked to explain this plant succession.