Variation of selfing rate and inbreeding depression among individuals and across generations within an admixed Cedrus population

[EN] We investigated the variation and short-term evolution of the selfing rate and inbreeding depression (ID) across three generations within a cedar forest that was established from admixture ca 1860. The mean selfing rate was 9.5%, ranging from 0 to 48% among 20 seed trees (estimated from paternally inherited chloroplast DNA). We computed the probability of selfing for each seed and we investigated ID by comparing selfed and outcrossed seeds within progenies, thus avoiding maternal effects. In all progenies, the germination rate was high (88-100%) and seedling mortality was low (0-12%). The germination dynamics differed significantly between selfed and outcrossed seeds within progenies in the founder gene pool but not in the following generations. This transient effect of selfing could be attributed to epistatic interactions in the original admixture. Regarding the seedling growth traits, the ID was low but significant: 8 and 6% for height and diameter growth, respectively. These rates did not vary among generations, suggesting minor gene effects. At this early stage, outcrossed seedlings outcompeted their selfed relatives, but not necessarily other selfed seedlings from other progenies. Thus, purging these slightly deleterious genes may only occur through within-family selection. Processes that maintain a high level of genetic diversity for fitness-related traits among progenies also reduce the efficiency of purging this part of the genetic load. © 2011 Macmillan Publishers Limited All rights reserved. Guardar / Salir Siguiente >This work has been partially supported by Grant PPI-00-04 from the Polytechnic University of Valencia (Spain). We thank B Fady and E Klein as well as two anonymous reviewers for their helpful comments on a previous version of the paper. We acknowledge B Jouaud, W Brunetto, F Jean and H Picot for seed collection and processing and laboratory assistance, as well as P Brahic and staff from the Experimental Nursery of Aix-Les Milles for nursery cares.Ferriol Molina, M.; Pichot, C.; Lefevre, F. (2011). Variation of selfing rate and inbreeding depression among individuals and across generations within an admixed Cedrus population. Heredity. 106(1):146-157. https://doi.org/10.1038/hdy.2010.451461571061Barret SH, Eckert CG (1990). Variation and evolution of mating systems in seed plants. In: Kawano S (ed). Biological Approaches and Evolutionary Trends in Plants. Academic Press: London. pp 230–254.Benton TG, Plaistow SJ, Coulson TN (2006). Complex population dynamics and complex causation: devils, details and demography. Proc R Soc B Biol Sci 273: 1173–1181.Bower AD, Aitken SN (2007). Mating system and inbreeding depression in whitebark pine (Pinus albicaulis Engelm.). Tree Genet Genomes 3: 379–388.Byers DL, Waller DM (1999). Do plant populations purge their genetic load? Effects of population size and mating history on inbreeding depression. Annu Rev Ecol Syst 30: 479–513.Cointat M (1996). Le roman du cèdre. Revue Forestière Française 48: 503–526.Collevatti RG, Grattapaglia D, Duvall J (2001). High resolution microsatellite based analysis of the mating system allows the detection of significant biparental inbreeding in Caryocar brasiliense, an endangered tropical tree species. Heredity 86: 60–67.Cottrell JE, White IMS (1995). The use of isozyme genetic markers to estimate the rate of outcrossing in a Sitka pruce (Picea sitchensis (Bong.) Carr.) seed orchard in Scotland. New Forests 10: 111–122.Coulson T, Benton TG, Lundberg P, Dall SRX, Kendall BE (2006). Putting evolutionary biology back in the ecological theatre: a demographic framework mapping genes to communities. Evol Ecol Res 8: 1155–1171.Durel CE, Bertin P, Kremer A (1996). Relationship between inbreeding depression and inbreeding coefficient in maritime pine (Pinus pinaster). Theor Appl Genet 92: 347–356.Eriksson E (2006). Thinning operations and their impact on biomass production in stands of Norway spruce and Scots pine. Biomass Bioenergy 30: 848–854.Fady B, Lefèvre F, Reynaud M, Vendramin GG, Bou Dagher-Karrat M, Anzidei M et al. (2003). Gene flow among different taxonomic units: evidence from nuclear and cytoplasmic markers in Cedrus plantation forests. Theor Appl Genet 107: 1132–1138.Farris MA, Mitton JB (1984). Population density, outcrossing rate, and heterozygote superiority in ponderosa pine. Evolution 38: 1151–1154.Favre-Duchartre M (1970). Des Ovules Aux Graines. Monographie 8. Masson et Cie.: Paris.Franklin EC (1969). Inbreeding Depression in Metrical Traits of Loblolly Pine (Pinus taeda L.) as a Result of Self-pollination. North Carolina State University: Raleigh, NC. Technical report No 40, School of Forest Resources.Gregorius HR, Ziehe M, Ross MD (1987). Selection caused by self-fertilization I. Four measures of self-fertilization and their effects on fitness. Theor Popul Biol 31: 91–115.Hamrick JL, Godt MJ (1989). Allozyme diversity in plant species. In: Brown AHD, Al Kahler MC, Weir BS (eds). Plant Population Genetics, Breeding, and Genetic Resources. Sinauer: Sunderland, MA. pp 43–63.Holsinger KE (1991). Mass-action models of plant mating systems—the evolutionary stability of mixed mating systems. Am Nat 138: 606–622.Husband BC, Schemske DW (1996). Evolution of the magnitude and timing of inbreeding depression in plants. Evolution 50: 54–70.Jones FA, Hamrick JL, Peterson CJ, Squiers ER (2006). Inferring colonization history from analyses of spatial genetic structure within populations of Pinus strobus and Quercus rubra. Mol Ecol 15: 851–861.Kärkkäinen K, Savolainen O (1993). The degree of early inbreeding depression determines the selfing rate at the seed stage: model and results from Pinus sylvestris (Scots pine). Heredity 71: 160–166.Keller LF, Waller DM (2002). Inbreeding effects in wild populations. Trends Ecol Evol 17: 230–241.Klein EK, Lavigne C, Gouyon PH (2006). Mixing of propagules from discrete sources at long distance: comparing an exponential tail to an exponential. BMC Ecol 6: 3.Knowles P, Furnier GR, Aleksiuk MK, Perry DJ (1987). Significant levels of self-fertilization in natural populations of tamarack. Can J Bot 65: 1087–1091.Koelewijn HP, Koski V, Savolainen O (1999). Magnitude and timing of inbreeding depression in Scots pine (Pinus sylvestris L.). Evolution 53: 758–768.Kremer A (1994). Genetic diversity and phenotypic variability of forest trees. Genet Sel Evol 26: s105–s123.Krouchi F, Derridj A, Lefèvre F (2004). Year and tree effect on reproductive organisation of Cedrus atlantica in a natural forest. For Ecol Manage 197: 181–189.Lande R (1988). Genetics and demography in biological conservation. Science 241: 1455–1460.Ledig FT (1986). Heterozygosity, heterosis, and fitness in outbreeding plants. In: Soulé ME (ed). Conservation Biology: the Science of Scarcity and Diversity. Sinauer Ass: Sunderland. pp 77–104.Lee JK, Nordheim EV, Kang H (1996). Inference for lethal gene estimation with application in plants. Biometrics 52: 451–462.Lefèvre F, Fady B, Fallour-Rubio D, Ghosn D, Bariteau M (2004). Impact of founder population, drift and selection on the genetic diversity of a recently translocated tree population. Heredity 93: 542–550.Marquardt PE, Epperson BK (2004). Spatial and population genetic structure of microsatellites in white pine. Mol Ecol 13: 3305–3315.Morgante M, Vendramin GG, Rossi P (1991). Effects of stand density on outcrossing rate in two Norway spruce (Picea abies) populations. Can J Bot 69: 2704–2708.Mosseler A, Major JE, Simpson JD, Daigle B, Lange K, Park YS et al. (2000). Indicators of population viability in red spruce, Picea rubens. I. Reproductive traits and fecundity. Can J Bot 78: 928–940.Naydenov KD, Tremblay FM, Alexandrov A, Fenton NJ (2005). Structure of Pinus sylvestris L. populations in Bulgaria revealed by chloroplast microsatellites and terpenes analysis : provenance tests. Biochem Syst Ecol 33: 1226–1245.Neale DB, Adams WT (1985). The mating system in natural and shelterwood stands of Douglas-fir. Theor Appl Genet 71: 201–207.Notivol E, Garcia-Gil MR, Alia R, Savolainen O (2007). Genetic variation of growth rhythm traits in the limits of a latitudinal cline in Scots pine. Can J For Res 37: 540–551.O’Connell LM, Russell J, Ritland K (2004). Fine-scale estimation of outcrossing in western redcedar with microsatellite assay of bulked DNA. Heredity 93: 443–449.Parducci L, Szmidt AE, Madaghiele A, Anzidei M, Vendramin GG (2001). Genetic variation at chloroplast microsatellites (CpSSRs) in Abies nebrodensis (Lojac.) Mattei and three neighboring Abies species. Theor Appl Genet 102: 733–740.Parraguirre-Lezama C, Vargas-Hernández JJ, Ramirez-Vallejo P, Ramirez Herrera C (2004). Mating system in four natural populations of Pinus greggii Engelm. Agrociencia 38: 107–119.Petit RJ, Hampe A (2006). Some evolutionary consequences of being a tree. Annu Rev Ecol Evol Syst 37: 187–214.Pichot C, Bastien C, Courbet F, Demesure-Musch B, Dreyfus P, Fady B et al. (2006). Déterminants et conséquences de la qualité génétique des graines et semis lors de la phase initiale de régénération naturelle des peuplements forestiers. In: 6e Colloque National du BRG ; La Rochelle 2006/10/02-04. Les Actes du Bureau des Ressources Génétiques 6: 277–297.Remington DL, O’Malley DM (2000a). Whole-genome characterization of embryonic stage inbreeding depression in a selfed loblolly pine family. Genetics 155: 337–348.Remington DL, O’Malley DM (2000b). Evaluation of major genetic loci contributing to inbreeding depression for survival and early growth in a selfed family of Pinus taeda. Evolution 54: 1580–1589.Restoux G, Silva DE, Sagnard F, Torre F, Klein E, Fady B (2008). Life at the margin: the mating system of Mediterranean conifers. Web Ecol 8: 94–102.Ribeiro MM, Mariette S, Vendramin GG, Szmidt AE, Plomion C, Kremer A (2002). Comparison of genetic diversity estimates within and among populations of maritime pine using chloroplast simple-sequence repeat and amplified fragment length polymorphism data. Mol Ecol 11: 869–877.Ritland K, El-Kassaby YA (1985). The nature of inbreeding in a seed orchard of Douglas fir as shown by an efficient multi-locus model. Theor Appl Genet 71: 375–384.Ritland K, Travis S (2004). Inferences involving individual coefficients of relatedness and inbreeding in natural populations of Abies. For Ecol Manage 197: 171–180.Robledo-Arnuncio JJ, Alia R, Gil L (2004). Increased selfing and correlated paternity in a small population of a predominantly outcrossing conifer, Pinus sylvestris. Mol Ecol 13: 2567–2577.Rouault G, Turgeon J, Candau JN, Roques A, Aderkas P (2004). Oviposition strategies of conifer seed chalcids in relation to host phenology. Naturwissenschaften 91: 472–480.Savolainen O, Kärkkäinen K, Kuittinen H (1992). Estimating numbers of embryonic lethals in conifers. Heredity 69: 308–314.Scofield DG, Schultz ST (2006). Mitosis, stature and evolution of plant mating systems: low-Phi and high-Phi plants. Proc R Soc B Biol Sci 273: 275–282.Shaw DV, Allard RW (1982). Estimation of outcrossing rates in douglas-fir using isoenzyme markers. Theor Appl Genet 62: 113–120.Skrøppa T (1996). Diallel crosses in Picea abies. II. Performance and inbreeding depression of selfed families. For Genet 3: 69–79.Sorensen FC (1997). Effects of sib mating and wind pollination on nursery seedling size, growth components, and phenology of Douglas-fir seed-orchard progenies. Can J For Res 27: 557–566.Sorensen FC (1999). Relationship between self-fertility, allocation of growth, and inbreeding depression in three coniferous species. Evolution 53: 417–425.Sorensen FC (2001). Effect of population outcrossing rate on inbreeding depression in Pinus contorta var. murrayana seedlings. Scand J For Res 16: 391–403.Sorensen FC, Adams WT (1993). Self fertility and natural selfing in three Oregon Cascade populations of lodgepole pine. In: Lindgren D (ed). Pinus contorta—From Untamed Forest to Domesticated Crop. Department of Forest Genetics and Plant Physiology, Sweden University of Agricultural Science: Umea, Sweden. Report 11, pp 358–374.Sorensen FC, Miles RS (1974). Self-pollination effects on Douglas fir and ponderosa pine seeds and seedlings. Silvae Genet 23: 135–138.Sorensen FC, Miles RS (1982). Inbreeding depression in height, height growth, and survival of Douglas-fir, ponderosa pine, and noble fir to 10 years of age. For Sci 28: 283–292.Terrab A, Paun O, Talavera S, Tremetsberger K, Arista M, Stuessy TF (2006). Genetic diversity and population structure in natural populations of Moroccan Atlas cedar (Cedrus atlantica; Pinaceae) determined with cpSSR markers. Am J Bot 93: 1274–1280.Vendramin GG, Lelli L, Rossi P, Morgante M (1996). A set of primers for the amplification of 20 chloroplast microsatellites in Pinaceae. Mol Ecol 5: 595–598.White TL, Adams WT, Neale DB (2007). Forest Genetics. CABI Publisher: Cambridge, MA. pp 149–186.Wilcox MD (1983). Inbreeding depression and genetic variances estimated from self- and cross- pollinated families of Pinus radiata. Silvae Genet 32: 89–96.Williams CG (2007). Re-thinking the embryo lethal system within the Pinaceae. Can J Bot 85: 667–677.Williams CG (2008). Selfed embryo death in Pinus taeda: a phenotypic profile. New Phytol 178: 210–222.Williams CG, Auckland LD, Reynolds MM, Leach KA (2003). Overdominant lethals as part of the conifer embryo lethal system. Heredity 91: 584–592.Wilson R (1923). Life history of Cedrus atlantica. Bot Gaz 75: 203–208.Yazdani R, Muona O, Rudin D, Szmidt AE (1985). Genetic structure of a Pinus sylvestris L. seed-tree stand and naturally regenerated understory. For Sci 31: 430–436

Accounting Problems Under the Excess Profits Tax

DNA vaccines based on subunits from pathogens have several advantages over other vaccine strategies. DNA vaccines can easily be modified, they show good safety profiles, are stable and inexpensive to produce, and the immune response can be focused to the antigen of interest. However, the immunogenicity of DNA vaccines which is generally quite low needs to be improved. Electroporation and co-delivery of genetically encoded immune adjuvants are two strategies aiming at increasing the efficacy of DNA vaccines. Here, we have examined whether targeting to antigen-presenting cells (APC) could increase the immune response to surface envelope glycoprotein (Env) gp120 from Human Immunodeficiency Virus type 1 (HIV- 1). To target APC, we utilized a homodimeric vaccine format denoted vaccibody, which enables covalent fusion of gp120 to molecules that can target APC. Two molecules were tested for their efficiency as targeting units: the antibody-derived single chain Fragment variable (scFv) specific for the major histocompatilibility complex (MHC) class II I-E molecules, and the CC chemokine ligand 3 (CCL3). The vaccines were delivered as DNA into muscle of mice with or without electroporation. Targeting of gp120 to MHC class II molecules induced antibodies that neutralized HIV-1 and that persisted for more than a year after one single immunization with electroporation. Targeting by CCL3 significantly increased the number of HIV-1 gp120-reactive CD8(+) T cells compared to non-targeted vaccines and gp120 delivered alone in the absence of electroporation. The data suggest that chemokines are promising molecular adjuvants because small amounts can attract immune cells and promote immune responses without advanced equipment such as electroporation.Funding Agencies|Research Council of Norway; Odd Fellow</p

Protective behaviour of citizens to transport accidents involving hazardous materials: A discrete choice experiment applied to populated areas nearby waterways

Background To improve the information for and preparation of citizens at risk to hazardous material transport accidents, a first important step is to determine how different characteristics of hazardous material transport accidents will influence citizens' protective behaviour. However, quantitative studies investigating citizens' protective behaviour in case of hazardous material transport accidents are scarce. Methods A discrete choice experiment was conducted among subjects (19-64 years) living in the direct vicinity of a large waterway. Scenarios were described by three transport accident characteristics: odour perception, smoke/vapour perception, and the proportion of people in the environment that were leaving at their own discretion. Subjects were asked to consider each scenario as realistic and to choose the alternative that was most appealing to them: staying, seekin

Association of cytokines with endothelium dependent flow mediated vasodilation (FMD) of systemic arteries in patients with non-ischemic cardiomyopathy

<p>Abstract</p> <p>Background</p> <p>Aim of this study was to elucidate the relation between localised inflammatory heart disease and endothelial dysfunction in the peripheral circulation, considering circulating cytokines as a potential link.</p> <p>Methods</p> <p>In 38 patients with non-ischemic heart disease, myocardial biopsies were examined for myocardial inflammation (immunohistology) and virus persistence (PCR). Cytokines (sIL-4, IFN-g, IFN-b, IFN-a, sIL-12p7, TNF-a) were measured by ELISA in venous serum. Endothelial function of the radial artery was examined by ultrasound, measuring diameter changes in response to reactive hyperemia (FMD), compared to glyceroltrinitrate (GTN-MD). Patients with EF < 35% were excluded.</p> <p>Results</p> <p>Age 44 ± 14 years, 19 male, 19 female, EF 63.5[16]%. FMD 4.38 [4.82]%. 30 patients had myocardial inflammation (8 not), 23 virus persistence (15 not). FMD correlated significantly with sIL-12p7 (p = 0.024, r = -0.365), but not with other cytokines. sIL-12p7 levels were significantly higher in patients with severely impaired FMD (n = 17), compared with normal FMD (n = 21): 10.70 [10.72] vs. 4.33 [7.81] pg/ml (p = 0.002). Endothelium independent vasodilation (GTN-MD 23.67 [8.21]%) was not impaired.</p> <p>Conclusion</p> <p>Endothelial dysfunction of peripheral arteries in patients with non-ischemic cardiomyopathy is associated with elevated serum concentrations of sIL-12p7, but not of other cytokines. Circulating sIL-12p7 may partly explain, that endothelial dysfunction is not restricted to the coronary circulation, but involves systemic arteries.</p

Rapidity and Centrality Dependence of Proton and Anti-proton Production from Au+Au Collisions at sqrt(sNN) = 130GeV

We report on the rapidity and centrality dependence of proton and anti-proton transverse mass distributions from Au+Au collisions at sqrt(sNN) = 130GeV as measured by the STAR experiment at RHIC. Our results are from the rapidity and transverse momentum range of |y|<0.5 and 0.35 <p_t<1.00GeV/c. For both protons and anti-protons, transverse mass distributions become more convex from peripheral to central collisions demonstrating characteristics of collective expansion. The measured rapidity distributions and the mean transverse momenta versus rapidity are flat within |y|<0.5. Comparisons of our data with results from model calculations indicate that in order to obtain a consistent picture of the proton(anti-proton) yields and transverse mass distributions the possibility of pre-hadronic collective expansion may have to be taken into account.Comment: 4 pages, 3 figures, 1 table, submitted to PR

Enhanced resistance to bacterial and fungal pathogens by overexpression of a human cathelicidin antimicrobial peptide (hCAP18/LL-37) in Chinese cabbage

The human cathelicidin antimicrobial protein hCAP18, which includes the C-terminal peptide LL-37, is a multifunctional protein. As a possible approach to enhancing the resistance to plant disease, a DNA fragment coding for hCAP18/LL-37 was fused at the C-terminal end of the leader sequence of endopolygalacturonase-inhibiting protein under the control of the cauliflower mosaic virus 35S promoter region. The construct was then introduced into Brassica rapa. LL-37 expression was confirmed in transgenic plants by reverse transcription-polymerase chain reaction and western blot analysis. Transgenic plants exhibited varying levels of resistance to bacterial and fungal pathogens. The average size of disease lesions in the transgenic plants was reduced to less than half of that in wild-type plants. Our results suggest that the antimicrobial LL-37 peptide is involved in wide-spectrum resistance to bacterial and fungal pathogen infection

Single Spin Asymmetry ANA_N in Polarized Proton-Proton Elastic Scattering at s=200\sqrt{s}=200 GeV

We report a high precision measurement of the transverse single spin asymmetry $A_N$ at the center of mass energy $\sqrt{s}=200$ GeV in elastic proton-proton scattering by the STAR experiment at RHIC. The $A_N$ was measured in the four-momentum transfer squared $t$ range $0.003 \leqslant |t| \leqslant 0.035$ \GeVcSq, the region of a significant interference between the electromagnetic and hadronic scattering amplitudes. The measured values of $A_N$ and its $t$-dependence are consistent with a vanishing hadronic spin-flip amplitude, thus providing strong constraints on the ratio of the single spin-flip to the non-flip amplitudes. Since the hadronic amplitude is dominated by the Pomeron amplitude at this $\sqrt{s}$, we conclude that this measurement addresses the question about the presence of a hadronic spin flip due to the Pomeron exchange in polarized proton-proton elastic scattering.Comment: 12 pages, 6 figure

Azimuthal anisotropy and correlations at large transverse momenta in p+pp+p and Au+Au collisions at sNN\sqrt{s_{_{NN}}}= 200 GeV

Results on high transverse momentum charged particle emission with respect to the reaction plane are presented for Au+Au collisions at $\sqrt{s_{_{NN}}}$= 200 GeV. Two- and four-particle correlations results are presented as well as a comparison of azimuthal correlations in Au+Au collisions to those in $p+p$ at the same energy. Elliptic anisotropy, $v_2$, is found to reach its maximum at $p_t \sim 3$ GeV/c, then decrease slowly and remain significant up to $p_t\approx 7$ -- 10 GeV/c. Stronger suppression is found in the back-to-back high-$p_t$ particle correlations for particles emitted out-of-plane compared to those emitted in-plane. The centrality dependence of $v_2$ at intermediate $p_t$ is compared to simple models based on jet quenching.Comment: 4 figures. Published version as PRL 93, 252301 (2004

Centrality and transverse momentum dependence of elliptic flow of multi-strange hadrons and ϕ\phi meson in Au+Au collisions at sNN\sqrt{s_{NN}} = 200 GeV

We present high precision measurements of elliptic flow near midrapidity ($|y|<1.0$) for multi-strange hadrons and $\phi$ meson as a function of centrality and transverse momentum in Au+Au collisions at center of mass energy $\sqrt{s_{NN}}=$ 200 GeV. We observe that the transverse momentum dependence of $\phi$ and $\Omega$ $v_{2}$ is similar to that of $\pi$ and $p$, respectively, which may indicate that the heavier strange quark flows as strongly as the lighter up and down quarks. This observation constitutes a clear piece of evidence for the development of partonic collectivity in heavy-ion collisions at the top RHIC energy. Number of constituent quark scaling is found to hold within statistical uncertainty for both 0-30$\%$ and 30-80$\%$ collision centrality. There is an indication of the breakdown of previously observed mass ordering between $\phi$ and proton $v_{2}$ at low transverse momentum in the 0-30$\%$ centrality range, possibly indicating late hadronic interactions affecting the proton $v_{2}$.Comment: 7 pages and 4 figures, Accepted for publication in Physical Review Letter