Evidence for adaptive radiation from a phylogenetic study of plant defenses.

One signature of adaptive radiation is a high level of trait change early during the diversification process and a plateau toward the end of the radiation. Although the study of the tempo of evolution has historically been the domain of paleontologists, recently developed phylogenetic tools allow for the rigorous examination of trait evolution in a tremendous diversity of organisms. Enemy-driven adaptive radiation was a key prediction of Ehrlich and Raven's coevolutionary hypothesis [Ehrlich PR, Raven PH (1964) Evolution 18:586-608], yet has remained largely untested. Here we examine patterns of trait evolution in 51 North American milkweed species (Asclepias), using maximum likelihood methods. We study 7 traits of the milkweeds, ranging from seed size and foliar physiological traits to defense traits (cardenolides, latex, and trichomes) previously shown to impact herbivores, including the monarch butterfly. We compare the fit of simple random-walk models of trait evolution to models that incorporate stabilizing selection (Ornstein-Ulenbeck process), as well as time-varying rates of trait evolution. Early bursts of trait evolution were implicated for 2 traits, while stabilizing selection was implicated for several others. We further modeled the relationship between trait change and species diversification while allowing rates of trait evolution to vary during the radiation. Species-rich lineages underwent a proportionately greater decline in latex and cardenolides relative to species-poor lineages, and the rate of trait change was most rapid early in the radiation. An interpretation of this result is that reduced investment in defensive traits accelerated diversification, and disproportionately so, early in the adaptive radiation of milkweeds.

Inbreeding and selection on sex ratio in the bark beetle Xylosandrus germanus.

ABSTRACT: BACKGROUND: Local Mate Competition (LMC) theory predicts a female should produce a more female-biased sex ratio if her sons compete with each other for mates. Because it provides quantitative predictions that can be experimentally tested, LMC is a textbook example of the predictive power of evolutionary theory. A limitation of many earlier studies in the field is that the population structure and mating system of the studied species are often estimated only indirectly. Here we use microsatellites to characterize the levels of inbreeding of the bark beetle Xylosandrus germanus, a species where the level of LMC is expected to be high. RESULTS: For three populations studied, genetic variation for our genetic markers was very low, indicative of an extremely high level of inbreeding (FIS = 0.88). There was also strong linkage disequilibrium between microsatellite loci and a very strong genetic differentiation between populations. The data suggest that matings among non-siblings are very rare (3%), although sex ratios from X. germanus in both the field and the laboratory have suggested more matings between non-sibs, and so less intense LMC. CONCLUSIONS: Our results confirm that caution is needed when inferring mating systems from sex ratio data, especially when a lack of biological detail means the use of overly simple forms of the model of interest.

Selection of common bean lines with high agronomic performance and high calcium and iron concentrations

The objective of this work was to evaluate the genetic variability of common bean lines for cycle, weight of 100 grains, grain yield, cooking time, and grain calcium and iron concentrations. Twenty-four common bean lines were evaluated in two crop cycles (2010 and 2011). The ¯Z index was used for the selection of superior lines for most of the traits. The DF 06-19, DF 06-03, DF 06-17, DF 06-20, DF 06-11, DF 06-14, DF 06-01, DF 06-08, DF 06-22, and DF 06-04 lines showed high grain yield. All lines were of semi-early cycle and of fast cooking. The DF 06-08 and DF 06-23 lines showed high calcium concentration in grains (>1.4 g kg-1 dry matter - DM), and the DF 06-09, DF 06-03, DF 06-04, and DF 06-06 lines presented high iron concentration in grains (>0.95 g kg-1 DM) in the two crop cycles. The DF 06-09 and DF 06-03 carioca lines present high agronomic performance and high iron concentration in grains. The DF 06-17 and DF 06-08 black lines present high agronomic performance and high calcium concentration in grains. The selection of the DF 06-09, DF 06-03, DF 06-17, and DF 06-08 lines is recommended.

Genetic parameters and simultaneous selection for root yield, adaptability and stability of cassava genotypes

The objective of this work was to estimate genetic parameters and to evaluate simultaneous selection for root yield and for adaptability and stability of cassava genotypes. The effects of genotypes were assumed as fixed and random, and the mixed model methodology (REML/Blup) was used to estimate genetic parameters and the harmonic mean of the relative performance of genotypic values (HMRPGV), for simultaneous selection purposes. Ten genotypes were analyzed in a complete randomized block design, with four replicates. The experiment was carried out in the municipalities of Altamira, Santarém, and Santa Luzia do Pará in the state of Pará, Brazil, in the growing seasons of 2009/2010, 2010/2011, and 2011/2012. Roots were harvested 12 months after planting, in all tested locations. Root yield had low coefficients of genotypic variation (4.25%) and broad-sense heritability of individual plots (0.0424), which resulted in low genetic gain. Due to the low genotypic correlation (0.15), genotype classification as to root yield varied according to the environment. Genotypes CPATU 060, CPATU 229, and CPATU 404 stood out as to their yield, adaptability, and stability.

The evolution of trans-generational altruism: kin selection meets niche construction.

A cornerstone result of sociobiology states that limited dispersal can induce kin competition to offset the kin selected benefits of altruism. Several mechanisms have been proposed to circumvent this dilemma but all assume that actors and recipients of altruism interact during the same time period. Here, this assumption is relaxed and a model is developed where individuals express an altruistic act, which results in posthumously helping relatives living in the future. The analysis of this model suggests that kin selected benefits can then feedback on the evolution of the trait in a way that promotes altruistic helping at high rates under limited dispersal. The decoupling of kin competition and kin selected benefits results from the fact that by helping relatives living in the future, an actor is helping individuals that are not in direct competition with itself. A direct consequence is that behaviours which actors gain by reducing the common good of present and future generations can be opposed by kin selection. The present model integrates niche-constructing traits with kin selection theory and delineates demographic and ecological conditions under which altruism can be selected for; and conditions where the 'tragedy of the commons' can be reduced.

Taajuusmuuttajan ohjelmiston testaus

Tämän diplomityön tavoitteena oli kehittää menetelmiä ja ohjeitataajuusmuuttajan sulautetun ohjelmiston kehityksen aikaiseen testaukseen. Soveltuvia menetelmiä etsittiin tutkimalla laajasti kirjallisuutta sekä selvittämälläyrityksen testauskäytäntöä. Tutkittuja kirjallisuudesta löytyneitä menetelmä olivat testauskehykset, simulointi ja staattinen sekä automaattinen testaus. Kirjallisuudesta etsittiin myös menetelmiä, joiden avulla testausprosessia voidaan helpottaa tai muuten parantaa. Tällaisista menetelmistä tutkittiin muun muassa testidatan valintaa, testauslähtöistä kehitystä sekä testattavuuden parantamista. Lisäksi selvitettiin uudelleenkäytettävien testien ohjelmointiin soveltuvia ohjelmointikieliä. Haastatteluiden ja dokumentaation avulla saatiin hyvä käsitys yrityksessä vallitsevasta testauskäytännöstä sekä sen ongelmakohdista. Testauksen ongelmiksi havaittiin testausprosessin järjestelmällisyyden puute sekä tarve suunnittelijoiden testauskoulutukseen. Testausprosessin parantamiseksi esitetään moduulitestauskehyksen käyttöönottoa. Lisäksi suunnittelijoiden testauskoulutuksella arvioidaan olevan suuri vaikutus koko testausprosessiin. Testitapausten suunnitteluun esitetään menetelmiä, joiden avulla voidaan suunnitella kattavampia testejä.

Early selection in open-pollinated Eucalyptus families based on competition covariates

The objetive of this work was to evaluate the influence of intergenotypic competition in open-pollinated families of Eucalyptus and its effects on early selection efficiency. Two experiments were carried out, in which the timber volume was evaluated at three ages, in a randomized complete block design. Data from the three years of evaluation (experiment 1, at 2, 4, and 7 years; and experiment 2, at 2, 5, and 7 years) were analyzed using mixed models. The following were estimated: variance components, genetic parameters, selection gains, effective number, early selection efficiency, selection gain per unit time, and coincidence of selection with and without the use of competition covariates. Competition effect was nonsignificant for ages under three years, and adjustment using competition covariates was unnecessary. Early selection for families is effective; families that have a late growth spurt are more vulnerable to competition, which markedly impairs ranking at the end of the cycle. Early selection is efficient according to all adopted criteria, and the age of around three years is the most recommended, given the high efficiency and accuracy rate in the indication of trees and families. The addition of competition covariates at the end of the cycle improves early selection efficiency for almost all studied criteria.

Selection of bee species for environmental risk assessment of GM cotton in the Brazilian Cerrado

The objective of this work was to list potential candidate bee species for environmental risk assessment (ERA) of genetically modified (GM) cotton and to identify the most suited bee species for this task, according to their abundance and geographical distribution. Field inventories of bee on cotton flowers were performed in the states of Bahia and Mato Grosso, and in Distrito Federal, Brazil. During a 344 hour sampling, 3,470 bees from 74 species were recovered, at eight sites. Apis mellifera dominated the bee assemblages at all sites. Sampling at two sites that received no insecticide application was sufficient to identify the three most common and geographically widespread wild species: Paratrigona lineata, Melissoptila cnecomola, and Trigona spinipes, which could be useful indicators of pollination services in the ERA. Indirect ordination of common wild species revealed that insecticides reduced the number of native bee species and that interannual variation in bee assemblages may be low. Accumulation curves of rare bee species did not saturate, as expected in tropical and megadiverse regions. Species-based approaches are limited to analyze negative impacts of GM cotton on pollinator biological diversity. The accumulation rate of rare bee species, however, may be useful for evaluating possible negative effects of GM cotton on bee diversity.

Carbon isotope fractionation for cotton genotype selection

The objective of this work was to evaluate the carbon isotope fractionation as a phenomic facility for cotton selection in contrasting environments and to assess its relationship with yield components. The experiments were carried out in a randomized block design, with four replicates, in the municipalities of Santa Helena de Goiás (SHGO) and Montividiu (MONT), in the state of Goiás, Brazil. The analysis of carbon isotope discrimination (Δ) was performed in 15 breeding lines and three cultivars. Subsequently, the root growth kinetic and root system architecture from the selected genotypes were determined. In both locations, Δ analyses were suitable to discriminate cotton genotypes. There was a positive correlation between Δ and seed-cotton yield in SHGO, where water deficit was more severe. In this site, the negative correlations found between Δ and fiber percentage indicate an integrative effect of gas exchange on Δ and its association with yield components. As for root robustness and growth kinetic, the GO 05 809 genotype performance contributes to sustain the highest values of Δ found in MONT, where edaphoclimatic conditions were more suitable for cotton. The use of Δ analysis as a phenomic facility can help to select cotton genotypes, in order to obtain plants with higher efficiency for gas exchange and water use.

Selection of the most informative morphoagronomic descriptors for cassava germplasm

The objective of this work was to select the most informative morphoagronomic descriptors for cassava (Manihot esculenta) germplasm and to evaluate the ability of different methods to select the descriptors. Ninety-five accessions were characterized using 51 morphoagronomic descriptors. Data were subjected to a multiple correspondence analysis (MCA), whose information was used in the following four methods of descriptor selection: reverse order of the descriptor for the pth factorial axis of the MCA (Jolliffe); sequential, multiple correspondence analysis (SMCA); mean of the contribution orders of the descriptor in the first three factorial axes (C3PA); and C3PA method weighted by the respective eigenvalues of the full analysis (C3PAWeig). The correlations between the dissimilarity matrix with all descriptors and the most informative descriptors were high and significant (0.75, 0.77, 0.83, and 0.84 for C3PAWeig, C3PA, SMCA, and Jolliffe, respectively). The less informative descriptors were discarded, considering those common among the selection methods and relevant for the breeding interests. Therefore, 32 morphoagronomic descriptors with correlation between the dissimilarity matrices (r=0.81) were selected, due to their high capacity to discriminate cassava germplasm and to their ability to maintain some preliminary agronomic traits, useful for the initial characterization of the germplasm.

Cuticular defects lead to full immunity to a major plant pathogen.

In addition to its role as a barrier, the cuticle is also a source of signals perceived by invading fungi. Cuticular breakdown products have been shown previously to be potent inducers of cutinase or developmental processes in fungal pathogens. Here the question was addressed as to whether plants themselves can perceive modifications of the cuticle. This was studied using Arabidopsis thaliana plants with altered cuticular structure. The expression of a cell wall-targeted fungal cutinase in A. thaliana was found to provide total immunity to Botrytis cinerea. The response observed in such cutinase-expressing plants is independent of signal transduction pathways involving salicylic acid, ethylene or jasmonic acid. It is accompanied by the release of a fungitoxic activity and increased expression of members of the lipid transfer protein, peroxidase and protein inhibitor gene families that provide resistance when overexpressed in wild-type plants. The same experiments were made in the bodyguard (bdg) mutant of A. thaliana. This mutant exhibits cuticular defects and remained free of symptoms after inoculation with B. cinerea. The expression of resistance was accompanied by the release of a fungitoxic activity and increased expression of the same genes as observed in cutinase-expressing plants. Structural defects of the cuticle can thus be converted into an effective multi-factorial defence, and reveal a hitherto hidden aspect of the innate immune response of plants.

The incidence and selection of multiple mating in plants.

Mating with more than one pollen donor, or polyandry, is common in land plants. In flowering plants, polyandry occurs when the pollen from different potential sires is distributed among the fruits of a single individual, or when pollen from more than one donor is deposited on the same stigma. Because polyandry typically leads to multiple paternity among or within fruits, it can be indirectly inferred on the basis of paternity analysis using molecular markers. A review of the literature indicates that polyandry is probably ubiquitous in plants except those that habitually self-fertilize, or that disperse their pollen in pollen packages, such as polyads or pollinia. Multiple mating may increase plants' female component by alleviating pollen limitation or by promoting competition among pollen grains from different potential sires. Accordingly, a number of traits have evolved that should promote polyandry at the flower level from the female's point of view, e.g. the prolongation of stigma receptivity or increases in stigma size. However, many floral traits, such as attractiveness, the physical manipulation of pollinators and pollen-dispensing mechanisms that lead to polyandrous pollination, have probably evolved in response to selection to promote male siring success in general, so that polyandry might often best be seen as a by-product of selection to enhance outcross siring success. In this sense, polyandry in plants is similar to geitonogamy (selfing caused by pollen transfer among flowers of the same plant), because both polyandry and geitonogamy probably result from selection to promote outcross siring success, although geitonogamy is almost always deleterious while polyandry in plants will seldom be so.

Pulmonary embolism severity index and troponin testing for the selection of low-risk patients with acute symptomatic pulmonary embolism.

Summary Background: The combination of the Pulmonary Embolism Severity Index (PESI) and troponin testing could help physicians identify appropriate patients with acute pulmonary embolism (PE) for early hospital discharge. Methods: This prospective cohort study included a total of 567 patients from a single center registry with objectively confirmed acute symptomatic PE. On the basis of the PESI, each patient was classified into 1 of 5 classes (I to V). At the time of hospital admission, patients had troponin I (cTnI) levels measured. The endpoint of the study was all-cause mortality within 30 days after diagnosis. We calculated the mortality rates in 4 patient groups: group 1: PESI class I-II plus cTnI <0.1 ng mL(-1); group 2: PESI classes III-V plus cTnI <0.1 ng mL(-1); group 3: PESI classes I-II plus cTnI >/= 0.1 ng mL(-1); and group 4: PESI classes III-V plus cTnI >/= 0.1 ng mL(-1). Results: The study cohort had a 30-day mortality of 10% (95% confidence interval [CI], 7.6 to 12.5%). Mortality rates in the 4 groups were 1.3%, 14.2%, 0% and 15.4%, respectively. Compared to non-elevated cTnl, the low-risk PESI had a higher negative predictive value (NPV) (98.9% vs 90.8%) and negative likelihood ratio (NLR) (0.1 vs 0.9) for predicting mortality. The addition of non-elevated cTnI to low-risk PESI did not improve the NPV or the NLR compared to either test alone. Conclusions: Compared to cTnl testing, PESI classification more accurately identified patients with PE who are at low risk of all-cause death within 30-days of presentation.

On the transcriptional role of NLRC5 and the function of NLRC5-driven MHC class I expression in the immune system

Major histocompatibility complex (MHC) molecules are of crucial importance for the immune system to recognize and defend the body against external attacks. Foreign antigens are presented by specialized cells, called antigen presenting cells, to T lymphocytes in the context of MHC molecules, thereby inducing T cell activation. In addition, MHC molecules are essential for Natural Killer (NK) cell biology, playing a role in NK cell education and activation. Recently, the NOD-like receptor (NLR) family member NLRC5 (NLR caspase recruitment domain containing protein 5) was found to act as transcriptional regulator of MHC class I, in particular in T and NK cells. Its role in MHC class I expression is however minor in dendritic cells (DCs). This raised the question of whether inflammatory conditions, which augment the levels of NLRC5 in DCs, could increase its contribution to MHC class I expression. Our work shows that MHC class I transcript and intracellular levels depend on NLRC5, while its role in MHC class I surface expression is instead negligible. We describe however a general salvage mechanism that enables cells with low intracellular MHC class I levels to nevertheless maintain relatively high MHC class I on the cell surface. In addition, we lack a thorough understanding of NLRC5 target gene specificity and mechanism of action. Our work delineates the unique consensus sequence in MHC class I promoters required for NLRC5 recruitment and pinpoints conserved features conferring its specificity. Furthermore, through genome-wide analyses, we confirm that NLRC5 regulates classical MHC class I genes and identify novel target genes all encoding non-classical MHC class I molecules exerting an array of functions in immunity and tolerance. We finally asked why a dedicated factor co-regulates MHC class I expression specifically in T and NK lymphocytes. We show that deregulated NLRC5 expression affects the education of NK cells and alters the crosstalk between T and NK cells, leading to NK cell-mediated killing of T lymphocytes. Altogether this thesis work brings insights into molecular and physiological aspects of NLRC5 function, which might help understand certain aspects of immune responses and disorders. -- Les molécules du complexe majeur d'histocompatibilité (CMH) sont essentielles au système immunitaire pour l'initiation de la réponse immunitaire. En effet, l'activation des lymphocytes T nécessite la reconnaissance d'un antigène étranger présenté par les cellules présentatrices d'antigènes sur une molécule du CMH. Les molécules du CMH ont également un rôle fondamental pour la fonction des cellules Natural Killer (NK) puisqu'elles sont nécessaires à leur processus d'éducation et d'activation. Récemment, NLRC5 (NLR caspase recruitment domain containing protein 5), un membre de la famille des récepteurs de type NOD (NLRs), a été décrit comme un facteur de transactivation de l'expression des gènes du CMH de classe I. A l'état basai, cette fonction transcriptionnelle est essentielle dans les lymphocytes T et NK, alors que ce rôle reste mineur pour l'expression des molécules du CMH de classe I dans les cellules dendritiques (DCs). Dans des conditions inflammatoires, l'expression de NLRC5 augmente dans les DCs. Notre travail démontre que, dans ces conditions, les transcrits et les niveaux intracellulaires des molécules du CMH de classe I augmentent aussi d'une façon dépendante de NLRC5. A contrario, le rôle de NLRC5 sur les niveaux de molécules de surface reste minoritaire. Cette observation nous a conduits à l'identification d'un mécanisme général de compensation qui permet aux cellules de maintenir des niveaux relativement élevés de molécules de CMH de class I à leur surface malgré de faibles niveaux intracellulaires. De plus, il semblait nécessaire de s'orienter vers une approche plus globale afin de déterminer l'étendue de la fonction transcriptionnelle de NLRC5. Par une approche du génome entier, nous avons pu décrire une séquence consensus conservée présente dans les promoteurs des gènes du CMH de classe I, sur laquelle NLRC5 est spécifiquement recruté. Nous avons pu également identifier de nouveaux gènes cibles codant pour des molécules de CMH de classe I non classiques impliqués dans l'immunité et la tolérance. Finalement, nous nous sommes demandé quel est l'intérêt d'avoir un facteur transcriptionnel, en l'occurrence NLRC5, qui orchestre l'expression du CMH de classe I dans les lymphocytes T et NK. Nous montrons que la dérégulation de l'expression de NLRC5 affecte l'éducation des cellules NK et conduit à la mort cellulaire des lymphocytes T médiée par les cellules NK. Dans l'ensemble ce travail de thèse contribue à la caractérisation du rôle de NLRC5, tant au niveau moléculaire que physiologique, ce qui présente un intérêt dans le cadre de la compréhension de certains aspects physiopathologique de la réponse immunitaire.

Market Area Selection of aSawmill Machinery Manufacturer

Tämän diplomityön tavoitteena on kartoittaa suomalaisen sahakonevalmistaja Veisto Oy:n kannalta lähitulevaisuuden merkittävimmät markkina-alueet, joiden sahateollisuuteen tehdään lähivuosina eniten korkean teknologian investointeja. Markkina-alueiden valinnassa sovelletaan sekä numeerisiin tilastoihin että asiantuntijahaastatteluihin pohjautuvia ranking-menetelmiä. Työn ensimmäinen osa käsittelee kansainvälisten teollisten markkinoiden ominaispiirteitä ja niiden analysointia. Pääpaino on kuitenkin screening-menetelmillä, markkina-alueiden vertailumenetelmilläja päätöksenteon työkaluilla. Työn toisessa osassa keskitytään markkina-alueiden screeningiin, analysointiin ja maiden eri ominaisuuksien vertailuun. Päätöksentekomatriiseja hyödyntäen valitaan Veisto Oy:lle kolme tällä hetkellä houkuttelevinta markkina-aluetta, joita ovat Venäjä, USA:n kaakkoisosan Southern Yellow Pine -alue sekä Etelä-Amerikan suurimmat sahaajamaat (Brasilia, Argentiina ja Chile) yhtenä alueena. Valituilla alueilla on Veiston kannalta omat haasteensa: USA:ssa vahvat kotimaiset kilpailijat ja uusien referenssien saaminen, Venäjällä investointien epävarmuus ja markkina-alueen laajuuden tuoma monimuotoisuus sekä Etelä-Amerikassa vahvat ruotsalaiset kilpailijat sekä etenkin Brasilian osalta tuntuvat suojatullit.