Yuval Sapir

Selection on domestication traits and quantitative trait loci in crop–wild sunflower hybrids

The strength and extent of gene flow from crops into wild populations depends, in part, on the fitness of the crop alleles, as well as that of alleles at linked loci. Interest in crop–wild gene flow has increased with the advent of transgenic plants, but nontransgenic crop–wild hybrids can provide case studies to understand the factors influencing introgression, provided that the genetic architecture and the fitness effects of loci are known. This study used recombinant inbred lines (RILs) generated from a cross between crop and wild sunflowers to assess selection on domestication traits and quantitative trait loci (QTL) in two contrasting environments, in Indiana and Nebraska, USA. Only a small fraction of plants (9%) produced seed in Nebraska, due to adverse weather conditions, while the majority of plants (79%) in Indiana reproduced. Phenotypic selection analysis found that a mixture of crop and wild traits were favoured in Indiana (i.e. had significant selection gradients), including larger leaves, increased floral longevity, larger disk diameter, reduced ray flower size and smaller achene (seed) mass. Selection favouring early flowering was detected in Nebraska. QTLs for fitness were found at the end of linkage groups six (LG6) and nine (LG9) in both field sites, each explaining 11–12% of the total variation. Crop alleles were favoured on LG9, but wild alleles were favoured on LG6. QTLs for numerous domestication traits overlapped with the fitness QTLs, including flowering date, achene mass, head number, and disk diameter. It remains to be seen if these QTL clusters are the product of multiple linked genes, or individual genes with pleiotropic effects. These results indicate that crop trait values and alleles may sometimes be favoured in a noncrop environment and across broad geographical regions.

Patterns of genetic and phenotypic variation in Iris haynei and I. atrofusca (Iris sect. Oncocyclus = the royal irises) along an ecogeographical gradient in Israel and the West Bank

Iris haynei and I. atrofusca are two closely related narrow endemics distributed vicariously along an ecogeographical north–south gradient in Israel and the West Bank. To obtain baseline information of the taxonomic status, conservation and population history of these taxa, we investigated patterns of phenotypic variation and the partitioning of genetic variation within and among populations using dominant random amplified polymorphic DNA (RAPD) markers. Multivariate (principal components analysis) and taxonomic distance analyses based on morphometric traits from eight populations revealed no unambiguous separation into two distinct groups. Results of genetic analyses for nine populations differed only slightly when either allele‐ or marker‐based approaches were employed. Mean within‐population diversity was high (0.258 for Nei’s expected heterozygosity), but there was no significant relationship between genetic diversity and either population size or latitude. Although the range‐wide estimate of GST (≈ 0.20) revealed relatively high differentiation among populations this value was inflated because of a small, but significant, component of molecular variance among regions viz. taxa (≈ 5%). Limited long‐distance dispersal capabilities in conjunction with a linearized habitat distribution are proposed to contribute to the approximate isolation by distance pattern observed. It also appears that extant populations are currently deviating from equilibrium conditions because of primary divergence of a formerly more widespread ancestral population. Given the absence of deep genetic and phenotypic subdivision among northern (I. haynei) vs. central/southern (I. atrofusca) populations, we argue for a revision of their species status. Nonetheless, we recommend conservation attention to these geographically differentiated segments as separate management units, which can be seen as an instructive example of incipient species formation.

The Genetic Architecture of Reproductive Isolation in Louisiana Irises: Pollination Syndromes and Pollinator Preferences

Abstract In animal-pollinated plants, pollinator preferences for divergent floral forms can lead to partial reproductive isolation. We describe regions of plant genomes that affect pollinator preferences for two species of Louisiana Irises, Iris brevicaulis and Iris fulva, and their artificial hybrids. Iris brevicaulis and I. fulva possess bee and bird-pollination syndromes, respectively. Hummingbirds preferred I. fulva and under-visited both I. brevicaulis and backcrosses toward this species. Lepidopterans preferred I. fulva and backcrosses toward I. fulva, but also under-visited I. brevicaulis and I. brevicaulis backcrosses. Bumblebees preferred I. brevicaulis and F1 hybrids and rarely visited I. fulva. Although all three pollen vectors preferred one or the other species, these preferences did not prevent visitation to other hybrid/parental classes. Quantitative trait locus (QTL) mapping, in reciprocal BC1 mapping populations, defined the genetic architecture of loci that affected pollinator behavior. We detected six and nine QTLs that affected pollinator visitation rates in the BCIb and BCIf mapping populations, respectively, with as many as three QTLs detected for each trait. Overall, this study reflects the possible role of quantitative genetic factors in determining (1) reproductive isolation, (2) the pattern of pollinator-mediated genetic exchange, and thus (3) hybrid zone evolution.

Genetic exchange and the origin of adaptations: prokaryotes to primates

Data supporting the occurrence of adaptive trait transfer (i.e. the transfer of genes and thus the phenotype of an adaptive trait through viral recombination, lateral gene transfer or introgressive hybridization) are provided in this review. Specifically, we discuss examples of lateral gene transfer and introgressive hybridization that have resulted in the transfer or de novo origin of adaptations. The evolutionary clades in which this process has been identified include all types of organisms. However, we restrict our discussion to bacteria, fungi, plants and animals. Each of these examples reflects the same consequence, namely that the transfer of genetic material, through whatever mechanism, may result in adaptive evolution. In particular, each of the events discussed has been inferred to impact adaptations to novel environmental settings in the recipient lineage.

Morning floral heat as a reward to the pollinators of the Oncocyclus irises

Relationships between flowering plants and their pollinators are usually affected by the amount of reward, mainly pollen or nectar, offered to pollinators by flowers, with these amounts usually positively correlated with floral display. The large Oncocyclus iris flowers, despite being the largest flowers in the East Mediterranean flora, are nectarless and have hidden pollen. No pollinators visit the flowers during daytime, and these flowers are pollinated only by night-sheltering solitary male bees. These iris flowers are partially or fully dark-colored, suggesting that they gather heat by absorbing solar radiation. Here we test the hypothesis that the dark-colored flowers of the Oncocyclus irises offer heat reward to their male solitary bee pollinators. Floral temperature was higher by 2.5°C than ambient air after sunrise. Solitary male bees emerged earlier after sheltering in Oncocyclus flowers than from other experimental shelter types. Pollination tunnels facing east towards the rising sun hosted more male bees than other aspects. We suggest that floral heat reward can explain the evolution of dark floral colors in Oncocyclus irises, mediated by the pollinators’ behavior.

Constructing Red Numbers for setting conservation priorities of endangered plant species: Israeli flora as a test case

Abstract A common problem in conservation policy is to define the priority of a certain species to invest conservation efforts when resources are limited. We suggest a method of constructing red numbers for plant species, in order to set priorities in conservation policy. The red number is an additive index, summarising values of four parameters: 1. Rarity – The number of sites (1 km2) where the species is present. A rare species is defined when present in 0.5% of the area or less. 2. Declining rate and habitat vulnerability – Evaluate the decreasing rate in the number of sites and/or the destruction probability of the habitat. 3. Attractivity – the flower size and the probability of cutting or exploitation of the plant. 4. Distribution type – scoring endemic species and peripheral populations. The plant species of Israel were scored for the parameters of the red number. Three hundred and seventy (370) species, 16.15% of the Israeli flora entered into the “Red List” received red numbers above 6. “Post mortem” analysis for the 34 extinct species of Israel revealed an average red number of 8.7, significantly higher than the average of the current red list. Only 15 species were known only from one site before extinction, indicating that rarity is not the only factor of extinction in Israeli flora. The red number suggested here is a pragmatic method and can be easily modified for conservation needs of any region. The red number method can supply a powerful quantitative weapon in the struggle for conservation.

Are pollinators the agents of selection for the extreme large size and dark color in Oncocyclus irises

Pollinator-mediated selection is a major evolutionary driver of floral traits; yet, such selection has rarely been tested for floral extreme traits. The Oncocyclus irises have exceptionally large, dark-colored flowers, associated with night-sheltering pollination and heat reward by the dark flowers. We quantified phenotypic selection on stem length, floral size and color in two species of iris (Iris atropurpurea and I. haynei), using an experimental approach. We estimated selection gradients for both flowers open to natural pollination and for flowers receiving supplementary hand pollination, assuming that open-pollinated flowers are affected by all factors that could influence fitness, whereas supplementary pollination removes the possible influence of pollinators. We found evidence for pollinator-mediated selection to increase floral size and stem length in I. atropurpurea, but floral color in this species was not under pollinator-mediated selection. In I. haynei, no pollinator-mediated selection on any of the traits was detected. We conclude that the extreme floral size of I. atropurpurea has probably evolved as a result of pollinator behavior. Lack of such evidence for I. haynei and for the dark floral color in both species suggests that other non-pollinator agents are selecting for these prominent traits, or that phenotypic color variation in these irises is neutral.

The endangered Iris atropurpurea (Iridaceae) in Israel: honey-bees, night-sheltering male bees and female solitary bees as pollinators.

BACKGROUND AND AIMS The coastal plain of Israel hosts the last few remaining populations of the endemic Iris atropurpurea (Iridaceae), a Red List species of high conservation priority. The flowers offer no nectar reward. Here the role of night-sheltering male solitary bees, honey-bees and female solitary bees as pollinators of I. atropurpurea is documented. METHODS Breeding system, floral longevity, stigma receptivity, visitation rates, pollen loads, pollen deposition and removal and fruit- and seed-set were investigated. KEY RESULTS The main wild pollinators of this plant are male eucerine bees, and to a lesser extent, but with the potential to transfer pollen, female solitary bees. Honey-bees were found to be frequent diurnal visitors; they removed large quantities of pollen and were as effective as male sheltering bees at pollinating this species. The low density of pollen carried by male solitary bees was attributed to grooming activities, pollen displacement when bees aggregated together in flowers and pollen depletion by honey-bees. In the population free of honey-bee hives, male bees carried significantly more pollen grains on their bodies. Results from pollen analysis and pollen deposited on stigmas suggest that inadequate pollination may be an important factor limiting fruit-set. In the presence of honey-bees, eucerine bees were low removal-low deposition pollinators, whereas honey-bees were high removal-low deposition pollinators, because they removed large amounts into corbiculae and deposited relatively little onto receptive stigmas. CONCLUSIONS Even though overall, both bee taxa were equally effective pollinators, we suggest that honey-bees have the potential to reduce the amount of pollen available for plant reproduction, and to reduce the amount of resources available to solitary bee communities. The results of this study have potential implications for the conservation of this highly endangered plant species if hives are permitted inside reserves, where the bulk of Oncocyclus iris species are protected.

Effects of floral traits and plant genetic composition on pollinator behavior

Pollinator-mediated selection plays a major role in floral evolution and speciation. Floral traits that influence animal pollinator behavior are the target of pollinator-mediated selection, but can only evolve if floral phenotypes have underlying genetic variation. Thus, understanding the genetic basis of a floral trait is a crucial step in studying pollinator-mediated selection. In this study I tested the effect of quantitative trait loci (QTL) underlying floral traits on pollinator behavior in recombinant inbred lines (RILs) in the common sunflower, Helianthus annuus L. and its crop relative. The indirect effects of QTL on pollinator behavior, mediated by floral phenotypes, were analyzed for six insect visitor types using structural equation modeling (SEM) and path analysis. For three of the six visitor types (large and small bees and non-bee insects) valid models were revealed when all three levels (QTL, floral traits, and pollinator behavior) were incorporated. Nested model without genetics were validated for five of the six visitor types. The results suggest that insect behavior as a reaction to floral phenotypes is affected by the genetic architecture of floral traits.

Species concepts and ecogeographical divergence of Oncocyclus irises

Several species concepts (SC) have been suggested as ways to view species, most of them based on Mayrs Biological SC. Recent research has raised the debate on the application of the species concepts to Oncocyclus irises. All the Oncocyclus species are able to produce vital progenies when crossed, thus, according to the Biological SC, they are a single biological species. Quantitative morphological characters are continuous among populations and also correlate with environmental conditions, preventing species delimitation according to Phenetic and Ecological SC. Genetic studies revealed high genetic diversity within the populations, thus, obscure delimitation by Phylogenetic SC. Pollination of the Oncocyclus irises is not species-specific, yet no transitional forms exist on the rare occasions of sympatric/parapatric distribution, roughly supporting delimitation according to the Recognition SC. The conclusion is that Oncocyclus populations do not clearly fit into species boundaries according to the relevan...