The evolution of unicoloniality in ant societies : recognition, dispersal and population genetic structure

Introduction Societies of ants, bees, wasps and termites dominate many terrestrial ecosystems (Wilson 1971). Their evolutionary and ecological success is based upon the regulation of internal conflicts (e.g. Ratnieks et al. 2006), control of diseases (e.g. Schmid-Hempel 1998) and individual skills and collective intelligence in resource acquisition, nest building and defence (e.g. Camazine 2001). Individuals in social species can pass on their genes not only directly trough their own offspring, but also indirectly by favouring the reproduction of relatives. The inclusive fitness theory of Hamilton (1963; 1964) provides a powerful explanation for the evolution of reproductive altruism and cooperation in groups with related individuals. The same theory also led to the realization that insect societies are subject to internal conflicts over reproduction. Relatedness of less-than-one is not sufficient to eliminate all incentive for individual selfishness. This would indeed require a relatedness of one, as found among cells of an organism (Hardin 1968; Keller 1999). The challenge for evolutionary biology is to understand how groups can prevent or reduce the selfish exploitation of resources by group members, and how societies with low relatedness are maintained. In social insects the evolutionary shift from single- to multiple queens colonies modified the relatedness structure, the dispersal, and the mode of colony founding (e.g. (Crozier & Pamilo 1996). In ants, the most common, and presumably ancestral mode of reproduction is the emission of winged males and females, which found a new colony independently after mating and dispersal flights (Hölldobler & Wilson 1990). The alternative reproductive tactic for ant queens in multiple-queen colonies (polygyne) is to seek to be re-accepted in their natal colonies, where they may remain as additional reproductives or subsequently disperse on foot with part of the colony (budding) (Bourke & Franks 1995; Crozier & Pamilo 1996; Hölldobler & Wilson 1990). Such ant colonies can contain up to several hundred reproductive queens with an even more numerous workforce (Cherix 1980; Cherix 1983). As a consequence in polygynous ants the relatedness among nestmates is very low, and workers raise brood of queens to which they are only distantly related (Crozier & Pamilo 1996; Queller & Strassmann 1998). Therefore workers could increase their inclusive fitness by preferentially caring for their closest relatives and discriminate against less related or foreign individuals (Keller 1997; Queller & Strassmann 2002; Tarpy et al. 2004). However, the bulk of the evidence suggests that social insects do not behave nepotistically, probably because of the costs entailed by decreased colony efficiency or discrimination errors (Keller 1997). Recently, the consensus that nepotistic behaviour does not occur in insect colonies was challenged by a study in the ant Formica fusca (Hannonen & Sundström 2003b) showing that the reproductive share of queens more closely related to workers increases during brood development. However, this pattern can be explained either by nepotism with workers preferentially rearing the brood of more closely related queens or intrinsic differences in the viability of eggs laid by queens. In the first chapter, we designed an experiment to disentangle nepotism and differences in brood viability. We tested if workers prefer to rear their kin when given the choice between highly related and unrelated brood in the ant F. exsecta. We also looked for differences in egg viability among queens and simulated if such differences in egg viability may mistakenly lead to the conclusion that workers behave nepotistically. The acceptance of queens in polygnous ants raises the question whether the varying degree of relatedness affects their share in reproduction. In such colonies workers should favour nestmate queens over foreign queens. Numerous studies have investigated reproductive skew and partitioning of reproduction among queens (Bourke et al. 1997; Fournier et al. 2004; Fournier & Keller 2001; Hammond et al. 2006; Hannonen & Sundström 2003a; Heinze et al. 2001; Kümmerli & Keller 2007; Langer et al. 2004; Pamilo & Seppä 1994; Ross 1988; Ross 1993; Rüppell et al. 2002), yet almost no information is available on whether differences among queens in their relatedness to other colony members affects their share in reproduction. Such data are necessary to compare the relative reproductive success of dispersing and non-dispersing individuals. Moreover, information on whether there is a difference in reproductive success between resident and dispersing queens is also important for our understanding of the genetic structure of ant colonies and the dynamics of within group conflicts. In chapter two, we created single-queen colonies and then introduced a foreign queens originating from another colony kept under similar conditions in order to estimate the rate of queen acceptance into foreign established colonies, and to quantify the reproductive share of resident and introduced queens. An increasing number of studies have investigated the discrimination ability between ant workers (e.g. Holzer et al. 2006; Pedersen et al. 2006), but few have addressed the recognition and discrimination behaviour of workers towards reproductive individuals entering colonies (Bennett 1988; Brown et al. 2003; Evans 1996; Fortelius et al. 1993; Kikuchi et al. 2007; Rosengren & Pamilo 1986; Stuart et al. 1993; Sundström 1997; Vásquez & Silverman in press). These studies are important, because accepting new queens will generally have a large impact on colony kin structure and inclusive fitness of workers (Heinze & Keller 2000). In chapter three, we examined whether resident workers reject young foreign queens that enter into their nest. We introduced mated queens into their natal nest, a foreign-female producing nest, or a foreign male-producing nest and measured their survival. In addition, we also introduced young virgin and mated queens into their natal nest to examine whether the mating status of the queens influences their survival and acceptance by workers. On top of polgyny, some ant species have evolved an extraordinary social organization called 'unicoloniality' (Hölldobler & Wilson 1977; Pedersen et al. 2006). In unicolonial ants, intercolony borders are absent and workers and queens mix among the physically separated nests, such that nests form one large supercolony. Super-colonies can become very large, so that direct cooperative interactions are impossible between individuals of distant nests. Unicoloniality is an evolutionary paradox and a potential problem for kin selection theory because the mixing of queens and workers between nests leads to extremely low relatedness among nestmates (Bourke & Franks 1995; Crozier & Pamilo 1996; Keller 1995). A better understanding of the evolution and maintenance of unicoloniality requests detailed information on the discrimination behavior, dispersal, population structure, and the scale of competition. Cryptic genetic population structure may provide important information on the relevant scale to be considered when measuring relatedness and the role of kin selection. Theoretical studies have shown that relatedness should be measured at the level of the `economic neighborhood', which is the scale at which intraspecific competition generally takes place (Griffin & West 2002; Kelly 1994; Queller 1994; Taylor 1992). In chapter four, we conducted alarge-scale study to determine whether the unicolonial ant Formica paralugubris forms populations that are organised in discrete supercolonies or whether there is a continuous gradation in the level of aggression that may correlate with genetic isolation by distance and/or spatial distance between nests. In chapter five, we investigated the fine-scale population structure in three populations of F. paralugubris. We have developed mitochondria) markers, which together with the nuclear markers allowed us to detect cryptic genetic clusters of nests, to obtain more precise information on the genetic differentiation within populations, and to separate male and female gene flow. These new data provide important information on the scale to be considered when measuring relatedness in native unicolonial populations.

Use of a dynamic foot pressure index to monitor the effects of treatment for equinus gait in children with cerebral palsy.

The purpose of this study is to introduce and describe a newly developed index using foot pressure analysis to quantify the degree of equinus gait in children with cerebral palsy before and after injection with botulinum toxin. Data were captured preinjection and 12 weeks postinjection. Ten children aged 2(1/2) to 6(1/2) years took part (5 boys and 5 girls). Three of them had a diagnosis of spastic diplegia and 7 of congenital hemiplegia. In total, 13 limbs were analyzed. After orientation and segmentation of raw pedobarographic data, we determined a dynamic foot pressure index graded 0 to 100 that quantified the relative degree of heel and forefoot contact during stance. These data were correlated (Pearson correlation) with clinical measurements of dorsiflexion at the ankle (on a slow and fast stretch) and video observation (using the Observational Gait Scale). Pedobarograph data were strongly correlated with both the Observational Gait Scale scores (R = 0.79, P < 0.005) and clinical measurements of dorsiflexion on a fast stretch, which is reflective of spasticity (R = 0.70, P < 0.005). We demonstrated the index's sensitivity in detecting changes in spasticity and good correlation with video observations seems to indicate this technique's potential validity. When manipulated and segmented appropriately, and with the development of a simple ordinal index, we found that foot pressure data provided a useful tool in tracking changes in patients with spastic equinus.

Empatía y tolerancia a la diversidad en un contexto educativo intercultural = Empathy and tolerance of diversity in an intercultural educative setting

Las relaciones entre empatía y conducta prosocial han estado ampliamente estudiadas desde hace años. Sin embargo, no existen estudios que utilicen estudiantes indígenas y mestizos de una universidad intercultural. El objetivo principal de la investigación fue analizar la tolerancia a la diversidad en relación a la empatía. La muestra estaba formada por 534 indígenas y mestizos, de edades comprendidas entre los 17 y los 22 años. Los resultados mostraron que los estudiantes con una alta capacidad empática eran también más tolerantes. Las chicas puntuaron significativamente superior en tolerancia y empatía que los chicos. Se encuentran diferencias entre indígenas y mestizos y entre universidad intercultural y universidad pública en relación a áreas específicas de la tolerancia a la diversidad

A simple and reliable method for the screening of transgenic tobacco plants

Even though much improvement has been made in plant transformation methods, the screening of transgenic plants is often a laborious work. Most approaches for detecting the transgene in transformed plants are still timeconsuming, and can be quite expensive. The objective of this study was to search for a simpler method to screen for transgenic plants. The infiltration of kanamycin (100 mg/mL) into tobacco leaves resulted in conspicuous chlorotic spots on the non-transgenic plant leaves, while no spots were seen on the leaves of transformed plants. This reaction occurred regardless of age of the tested plants, and the method has proven to be simple, fast, non-destructive, relatively cheap, and reliable. These results were comparable to those obtained by the polymerase chain reaction (PCR) amplification of the transgene using specific primers.

Isozymic variability in a Brazilian collection of annatto (Bixa orellana L.)

The objectives of this work were to optimize the isozyme electrophoresis technique for Bixa orellana, and use isozyme markers for a preliminary survey on the genetic variability in Brazilian annatto germplasm accessions. Collection consisted of seed samples from sixty open pollinated trees, representing two Northern and four Southern geographic provenances. The extraction, electrophoresis, and interpretation of annatto isozymes are described. Three out of the twenty-one identified isozyme loci were polymorphic in the collection. The percentage of polymorphic loci (P = 21.05) and the expected heterozygosity in annatto (H T = 0.064) were low, compared to other tropical woody species. A UPGMA phenogram, constructed with Nei's genetic distances, clearly separated the germplasm provenant from North and Central Brazil. Variability was significantly higher among the accessions from Maranhão. A sharp genetic differentiation was detected between accessions from Maranhão and Pará States, despite their geographical proximity. The distinctive isozyme polymorphism, observed in the accessions from Maranhão, together with reports on local morphological heterogeneity in annatto fruit shape, color, and pubescence, calls for more detailed genetic and taxonomic investigation.

Alho em pó na alimentação alternativa de frangos de corte

O objetivo deste trabalho foi estudar o efeito da inclusão de alho em pó (Allium sativum) em rações para frangos, sobre desempenho, rendimento de carcaça e partes, peso de órgãos, níveis de colesterol e triacilgliceróis sangüíneos, e morfometria intestinal. Foram utilizados 720 pintainhos machos Cobb, com um dia de idade, distribuídos em blocos casualizados com seis tratamentos: ração sem promotor de crescimento (PC), sem anticoccidiano (AC) e com 0,00, 0,25, 0,50, 0,75 e 1,00% de alho; e ração com PC+AC, com quatro repetições de 30 aves cada. Peso, ganho de peso, consumo de ração, conversão alimentar e mortalidade foram obtidos aos 21 e 42 dias de idade. Aos 42 dias de idade foram abatidas 20 aves por tratamento para determinação do rendimento de carcaça e 4 aves para avaliação do peso de órgãos e morfometria intestinal. Sangue de 8 aves por tratamento foi colhido para determinação de colesterol e triacilgliceróis. Em aves alimentadas até 42 dias de idade com ração sem promotor de crescimento e anticoccidiano, a inclusão de até 1,00% de alho em pó beneficiou a conversão alimentar e não alterou o rendimento de carcaça e partes, peso relativo dos órgãos, mucosa intestinal, nem os níveis séricos de colesterol e triacilgliceróis. O alho não substitui com eficiência o antibiótico usado como promotor de crescimento em rações de frangos.

Silagem de grãos úmidos de milho em substituição ao milho seco da ração de frangos de corte criados em dois sistemas

O objetivo do trabalho foi avaliar a inclusão de silagem de grãos úmidos de milho, em substituição ao milho seco, em rações para frangos de corte criados nos sistemas convencional e alternativo. Foram utilizados 720 pintos machos Cobb, com um dia de idade, distribuídos em blocos ao acaso, em esquema fatorial 2x3 (dois sistemas de criação: convencional e alternativo; três níveis de silagem de grãos úmidos, em substituição ao milho seco: 0%, 30% e 60%), com quatro repetições cada. Foram obtidas médias de peso, ganho de peso, consumo de ração, conversão alimentar, fator de produção e mortalidade aos 21, 42 e 49 dias de idades. Aos 49 dias, foram abatidas 28 aves, por tratamento, para determinação do rendimento de carcaça, cortes e gordura abdominal. A inclusão de até 60% de silagem de grãos úmidos, em substituição ao milho seco, reduziu o custo da ração, sem alterar o desempenho das aves aos 21, 42 e 49 dias, nem o rendimento de carcaça aos 49 dias. No sistema de criação alternativo, independentemente do nível de inclusão de silagem de grãos úmidos, o desempenho dos frangos foi inferior, porém o retorno econômico foi maior.

Energia metabolizável de alimentos na formulação de ração para frangos de corte

O objetivo deste trabalho foi avaliar o efeito da formulação de rações para frangos de corte, com a utilização de valores de energia metabolizável (EM) dos alimentos determinados por diferentes métodos. O delineamento experimental foi inteiramente casualizado, com quatro tratamentos, que consistiram na formulação de rações baseadas na: EM aparente corrigida e EM aparente, ambas determinadas pelo método da coleta total com pintos; EM aparente corrigida, determinada pelo método da coleta total com galos; e EM verdadeira corrigida, determinada pelo método da alimentação forçada com galos. Na fase inicial, foi obtido maior consumo de ração, ganho de peso e melhor conversão alimentar com a formulação baseada na EM aparente corrigida (pintos). Durante a fase final, a formulação com EM verdadeira corrigida (galos) promoveu maior ganho de peso e melhor conversão alimentar. No período de 1 a 49 dias, as aves alimentadas com as rações formuladas com EM aparente corrigida (pintos) tiveram melhor desempenho. As características de carcaça não foram influenciadas pelos tratamentos. Até os 21 dias de idade, deve-se considerar os valores de EM aparente corrigida (pintos), para a formulação das rações de frangos de corte; após essa idade, deve-se considerar os valores de EM aparente corrigida (galos) ou EM verdadeira corrigida (galos).

Características de carcaça de cordeiros em pastagem de azevém manejada em diferentes alturas

O objetivo deste trabalho foi avaliar as características de carcaça, de cordeiros mantidos em pastagem de azevém anual (Lolium multiflorum Lam.) manejada em diferentes alturas (5, 10, 15 e 20 cm), sob lotação contínua com carga variável, no período de agosto a novembro de 1999. Utilizou-se um delineamento em blocos ao acaso, com três repetições, e as variáveis estudadas foram submetidas à análise de regressão. Os pesos de abate, carcaça quente, filé mignon, paleta e costela diferiram entre os tratamentos e foram inferiores no tratamento relativo à altura de manejo de 5 cm, quando comparados aos tratamentos de 10, 15 e 20 cm, que não diferiram entre si. Não houve diferença entre tratamentos para os pesos de pernil e de carré. O rendimento de carcaça variou entre 42,8±0,29 e 44,1±0,32%. Todas as variáveis apresentaram relação quadrática com os tratamentos. O ganho médio diário influenciou diretamente o peso de abate que, por sua vez, correlacionou fortemente com o rendimento e o peso da carcaça. Os resultados mostram que os melhores rendimentos dos cortes comerciais de cordeiros são obtidos quando a pastagem de azevém é manejada no intervalo entre 12 e 14 cm de altura.

GWAS of 126,559 individuals identifies genetic variants associated with educational attainment.

A genome-wide association study (GWAS) of educational attainment was conducted in a discovery sample of 101,069 individuals and a replication sample of 25,490. Three independent single-nucleotide polymorphisms (SNPs) are genome-wide significant (rs9320913, rs11584700, rs4851266), and all three replicate. Estimated effects sizes are small (coefficient of determination R(2) ≈ 0.02%), approximately 1 month of schooling per allele. A linear polygenic score from all measured SNPs accounts for ≈2% of the variance in both educational attainment and cognitive function. Genes in the region of the loci have previously been associated with health, cognitive, and central nervous system phenotypes, and bioinformatics analyses suggest the involvement of the anterior caudate nucleus. These findings provide promising candidate SNPs for follow-up work, and our effect size estimates can anchor power analyses in social-science genetics.

Manejo da pastagem de azevém, contaminação larval no pasto e infecção parasitária em ovinos

O objetivo deste trabalho foi avaliar o efeito do manejo da pastagem de azevém sobre o risco de reinfecção parasitária. Foram utilizadas duas ofertas (10 e 20 kg de matéria seca por 100 kg de peso vivo) em dois métodos de pastejo (lotação contínua e intermitente). A infestação parasitária no mantilho e nos diferentes estratos da forragem foi avaliada nos estratos: acima de 15, 10-15, 5-10, 2,5-5 e 0-2,5 cm; a carga parasitária no animal foi avaliada por exames coproparasitológicos; e a seletividade da dieta foi avaliada com afilhos marcados. Independentemente do método, a densidade larval aumentou do topo para a base do dossel forrageiro. No entanto, entre os estratos aptos ao pastejo foram observadas diferenças na oferta de 20% do peso vivo. A oferta influenciou o número de larvas recuperadas na forragem e a infecção no animal para ambos os gêneros de parasitas Haemonchus spp. e Trichostrongylus spp. A maior oferta proporcionou maior contaminação larval e infecção nos ovinos, porém na oferta de 10% do peso vivo, a quantidade de larvas presentes no pasto foi menor em lotação contínua, mas a carga parasitária no animal foi semelhante à da lotação rotacionada, independentemente do método de pastejo.

The genome sequence of taurine cattle: a window to ruminant biology and evolution.

To understand the biology and evolution of ruminants, the cattle genome was sequenced to about sevenfold coverage. The cattle genome contains a minimum of 22,000 genes, with a core set of 14,345 orthologs shared among seven mammalian species of which 1217 are absent or undetected in noneutherian (marsupial or monotreme) genomes. Cattle-specific evolutionary breakpoint regions in chromosomes have a higher density of segmental duplications, enrichment of repetitive elements, and species-specific variations in genes associated with lactation and immune responsiveness. Genes involved in metabolism are generally highly conserved, although five metabolic genes are deleted or extensively diverged from their human orthologs. The cattle genome sequence thus provides a resource for understanding mammalian evolution and accelerating livestock genetic improvement for milk and meat production.

Association analyses of 249,796 individuals reveal 18 new loci associated with body mass index.

Obesity is globally prevalent and highly heritable, but its underlying genetic factors remain largely elusive. To identify genetic loci for obesity susceptibility, we examined associations between body mass index and ∼ 2.8 million SNPs in up to 123,865 individuals with targeted follow up of 42 SNPs in up to 125,931 additional individuals. We confirmed 14 known obesity susceptibility loci and identified 18 new loci associated with body mass index (P < 5 × 10⁻⁸), one of which includes a copy number variant near GPRC5B. Some loci (at MC4R, POMC, SH2B1 and BDNF) map near key hypothalamic regulators of energy balance, and one of these loci is near GIPR, an incretin receptor. Furthermore, genes in other newly associated loci may provide new insights into human body weight regulation.

Genetic studies of body mass index yield new insights for obesity biology

Obesity is heritable and predisposes to many diseases. To understand the genetic basis of obesity better, here we conduct a genome-wide association study and Metabochip meta-analysis of body mass index (BMI), a measure commonly used to define obesity and assess adiposity, in up to 339,224 individuals. This analysis identifies 97 BMI-associated loci (P < 5 × 10(-8)), 56 of which are novel. Five loci demonstrate clear evidence of several independent association signals, and many loci have significant effects on other metabolic phenotypes. The 97 loci account for ∼2.7% of BMI variation, and genome-wide estimates suggest that common variation accounts for >20% of BMI variation. Pathway analyses provide strong support for a role of the central nervous system in obesity susceptibility and implicate new genes and pathways, including those related to synaptic function, glutamate signalling, insulin secretion/action, energy metabolism, lipid biology and adipogenesis.