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Distyly is an iconic floral polymorphism governed by a supergene, which promotes efficient pollen transfer and outcrossing through reciprocal differences in the position of sexual organs in flowers, often coupled with heteromorphic self-incompatibility. Distyly has evolved convergently in multiple flowering plant lineages, but has also broken down repeatedly, often resulting in homostylous, self-compatible populations with elevated rates of self-fertilization. Here, we aimed to study the genetic causes and genomic consequences of the shift to homostyly in Linum trigynum, which is closely related to distylous Linum tenue. Building on a high-quality genome assembly, we show that L. trigynum harbors a genomic region homologous to the dominant haplotype of the distyly supergene conferring long stamens and short styles in L. tenue, suggesting that loss of distyly first occurred in a short-styled individual. In contrast to homostylous Primula and Fagopyrum, L. trigynum harbors no fixed loss-of-function mutations in coding sequences of S-linked distyly candidate genes. Instead, floral gene expression analyses and controlled crosses suggest that mutations downregulating the S-linked LtWDR-44 candidate gene for male self-incompatibility and/or anther height could underlie homostyly and self-compatibility in L. trigynum. Population genomic analyses of 224 whole-genome sequences further demonstrate that L. trigynum is highly self-fertilizing, exhibits significantly lower genetic diversity genome-wide, and is experiencing relaxed purifying selection and less frequent positive selection on nonsynonymous mutations relative to L. tenue. Our analyses shed light on the loss of distyly in L. trigynum, and advance our understanding of a common evolutionary transition in flowering plants.
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Flores , Genoma de Planta , Flores/genéticaRESUMEN
Supergenes govern multi-trait-balanced polymorphisms in a wide range of systems; however, our understanding of their origins and evolution remains incomplete. The reciprocal placement of stigmas and anthers in pin and thrum floral morphs of distylous species constitutes an iconic example of a balanced polymorphism governed by a supergene, the distyly S-locus. Recent studies have shown that the Primula and Turnera distyly supergenes are both hemizygous in thrums, but it remains unknown whether hemizygosity is pervasive among distyly S-loci. As hemizygosity has major consequences for supergene evolution and loss, clarifying whether this genetic architecture is shared among distylous species is critical. Here, we have characterized the genetic architecture and evolution of the distyly supergene in Linum by generating a chromosome-level genome assembly of Linum tenue, followed by the identification of the S-locus using population genomic data. We show that hemizygosity and thrum-specific expression of S-linked genes, including a pistil-expressed candidate gene for style length, are major features of the Linum S-locus. Structural variation is likely instrumental for recombination suppression, and although the non-recombining dominant haplotype has accumulated transposable elements, S-linked genes are not under relaxed purifying selection. Our findings reveal remarkable convergence in the genetic architecture and evolution of independently derived distyly supergenes, provide a counterexample to classic inversion-based supergenes, and shed new light on the origin and maintenance of an iconic floral polymorphism.
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Lino , Lino/genética , Elementos Transponibles de ADN , Flores/genética , Genómica , Sitios Genéticos , Evolución MolecularRESUMEN
Supergenes are genomic regions containing sets of tightly linked loci that control multi-trait phenotypic polymorphisms under balancing selection. Recent advances in genomics have uncovered significant variation in both the genomic architecture as well as the mode of origin of supergenes across diverse organismal systems. Although the role of genomic architecture for the origin of supergenes has been much discussed, differences in the genomic architecture also subsequently affect the evolutionary trajectory of supergenes and the rate of degeneration of supergene haplotypes. In this review, we synthesize recent genomic work and historical models of supergene evolution, highlighting how the genomic architecture of supergenes affects their evolutionary fate. We discuss how recent findings on classic supergenes involved in governing ant colony social form, mimicry in butterflies, and heterostyly in flowering plants relate to theoretical expectations. Furthermore, we use forward simulations to demonstrate that differences in genomic architecture affect the degeneration of supergenes. Finally, we discuss implications of the evolution of supergene haplotypes for the long-term fate of balanced polymorphisms governed by supergenes.
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Evolución Molecular , Insectos/genética , Animales , Evolución Biológica , Mimetismo Biológico , Estructuras Genéticas , Haplotipos , Polimorfismo Genético , Recombinación GenéticaRESUMEN
Plants adapted to extreme conditions can be at high risk from climate change; arctic-alpine plants, in particular, could "run out of space" as they are out-competed by expansion of woody vegetation. Mountain regions could potentially provide safe sites for arctic-alpine plants in a warmer climate, but empirical evidence is fragmentary. Here we present a 24,000-year record of species persistence based on sedimentary ancient DNA (sedaDNA) from Lake Bolshoye Shchuchye (Polar Urals). We provide robust evidence of long-term persistence of arctic-alpine plants through large-magnitude climate changes but document a decline in their diversity during a past expansion of woody vegetation. Nevertheless, most of the plants that were present during the last glacial interval, including all of the arctic-alpines, are still found in the region today. This underlines the conservation significance of mountain landscapes via their provision of a range of habitats that confer resilience to climate change, particularly for arctic-alpine taxa.
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Cambio Climático , Ecosistema , Desarrollo de la Planta , Plantas/clasificación , Regiones ÁrticasRESUMEN
BACKGROUND: Early life stress is a known risk factor for diseases and premature death. We tested whether parenting style impacts telomere length (TL), a cellular aging biomarker. METHODS: Information on parents' style of parenting was obtained from 199 participants in the Adventist Health Study-1 (AHS-1) who 27+ years later also enrolled in the AHS-2 where blood was collected for relative TL (rTL) assessment. RESULTS: Subjects describing their mothers' parenting style as cold had on average 25% smaller rTL compared to subjects not reporting a cold mother (1.89 vs 2.53). This association was greatest among those with less education, and those who stayed overweight/obese or put on weight during follow-up. CONCLUSIONS: These results support previous findings that early life stress may have health implications by promoting cellular aging, and expands these stressors to include cold parenting during an individuals' formative years. Higher education and normal weight seem to provide some resilience.
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Senescencia Celular/fisiología , Responsabilidad Parental/psicología , Padres/psicología , Estrés Psicológico/fisiopatología , Estrés Psicológico/psicología , Adulto , Femenino , Humanos , Masculino , Madres/psicología , Obesidad/fisiopatología , Obesidad/psicología , Estudios Retrospectivos , Factores de Riesgo , TelómeroRESUMEN
Magnetic perturbation measurements will be invaluable for characterizing Lithium Tokamak Experiment Beta (LTX-ß) plasmas due to the time-evolving 3D nature of the magnetic fields generated by eddy currents in the vessel and copper shell segments, as well as enhanced MHD instability drive due to newly introduced neutral beam heating. The LTX-ß upgrade includes two new arrays of Mirnov coils: a shell eddy sensor array of two-axis coils distributed over the back surface of one shell segment and a toroidal array of poloidal field coils at the low-field side midplane gap. Evaporative lithium wall-coating and the high temperatures required for liquid lithium wall operation both complicate the implementation of in-vessel diagnostics. While the shell array is protected from lithium exposure, the shell segment to which it is mounted will at times exceed 300 °C. The toroidal array, however, will experience direct line-of-sight exposure to the lithium evaporator as well as close proximity to the hot shell and may also be subject to poorly confined beam-driven fast ions. We describe how the two new Mirnov coil arrays meet these environmental challenges and enhance the LTX-ß diagnostic suite.
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Seed heteromorphism is the reproductive strategy characterized by the simultaneous production of multiple seed types. While comparing heteromorphic to monomorphic strategies is mathematically simple, there is no explicit test for assessing which ratio of seed morphs minimizes fitness variance, and hence offers a basis for comparing different heteromorphic strategies. Such a test may be particularly valuable when more than two distinct morphs are present, since many strategies may have equivalent geometric fitnesses. As noted by Gillespie (1974), in these cases avoiding rare but evolutionarily important instances of severe reductions in fitness involves the minimization of variation in fitness-i.e., risk. Here I compute the optimal proportions of two or more seed morphs for heteromorphic strategies that either: (1) minimize total fitness variance; or (2) maximize the fitness-risk ratio-i.e., the "extra" fitness accrued per unit of "extra" fitness variance. This work thereby provides a testable null hypothesis to estimate the optimal frequencies of seed morphs when multiple heteromorphic strategies have evolved in environments with severe fitness risks. Moreover, it also permits the calculation of expected seed morph frequencies when more than two seed morphs are produced.
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A mixture of proprietary anthelmintics delivering 0.5 mg/kg moxidectin, 9.06 mg/kg oxfendazole, 15 mg/kg levamisole, and 0.08 mg/kg selenium on bodyweight basis per os to red deer is investigated. On a deer farm with a history of parasite problems, six weaner red deer were treated orally with a 50/50 mixture of Exodus Pour-On and Oxfen C Plus (Ex/Ox) at a dose rate of 1 ml/5 kg bodyweight. Six herd mates were untreated. Eleven days later abomasal worm counts for the untreated deer revealed an arithmetic mean burden of 2,566 Ostertagia-type worms and 300 Trichostrongylus axei. No worms were detected in the abomasa of the treated group. Six yearling red deer were treated with the Ex/Ox combination and sent 39 days later to a slaughter plant where tissue samples were collected for residue analysis. Moxidectin was the only anthelmintic compound to show residues and the concentrations measured were well below maximum residue limits. Laboratory analysis of the Ex/Ox product after six-week storage at ambient temperature indicated good physical and chemical stability. These investigations support the hypothesis that the Ex/Ox combination can be an effective and practical anthelmintic option for use in red deer against a background of widespread gastrointestinal parasite resistance to the registered alternatives.
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Detailed investigation of landforms and their underlying deposits on the eastern margin of Fenland, East Anglia, demonstrated that they represent a series of glaciofluvial delta-fan and related sediments. Associated with these deposits are glacially dislocated sediments including tills, meltwater and pre-existing fluvial sediments. These 'Skertchly Line' deposits occur in the context of a substantial ice lobe that entered Fenland from the N to NE, dammed the streams entering the basin and caused glacial lakes to form in the valleys on the margins. Bulldozing by the ice lobe caused a series of ice-pushed ridges to form at the dynamic margin, especially at the ice maximum and during its retreat phases. Meltwater formed a series of marginal fans that coalesced into marginal accumulations in the SE of the basin. The ice lobe is named the Tottenhill glaciation. Further investigations of the Fenland margin have revealed the extent of the Tottenhill glaciation in the Fenland Basin, to the south and west, in sufficient detail to demonstrate the nature of the Tottenhill ice lobe and the landscape left on deglaciation. The ice lobe is likely to have been prone to surging. This is indicated by the low gradient of the ice lobe, the presence of underlying ductile Mesozoic clays, the evidence of ice-marginal flooding and the presence of arcuate glaciotectonic push moraines. Regional correlation, supported by independent numerical geochronology, indicates that the glaciation occurred ca 160 ka, i.e. during the late Middle Pleistocene, Marine Isotope Stage (MIS) 6, the Wolstonian Stage. Comparison and correlation across the southern North Sea Basin confirms that the glaciation is the equivalent of that during the Late Saalian Drenthe Stadial in The Netherlands. The implications of this correlation are presented. Before the glaciation occurred, the Fenland Basin did not exist. It appears to have been initiated by a subglacial tunnel valley system beneath the Anglian (=Elsterian, MIS 12) ice sheet. During the subsequent Hoxnian (=Holsteinian; approx. MIS 11) interglacial, the sea invaded the drainage system inherited following the glacial retreat. The evolution through the subsequent ca 200 ka Early to Middle Wolstonian substages, the interval between the Hoxnian (Holsteinian) temperate Stage and the Wolstonian glaciation, represents a period during which fluvial and periglacial activity modified the landscape under cold climates, and organic sediments were laid down during a warmer event. Palaeolithic humans were also periodically present during this interval, their artefacts having been reworked by the subsequent glaciation. The deglaciation was followed by re-establishment of the rivers associated with the deposition of Late Wolstonian (Warthe Stadial) gravels and sands, and later, deposits of the Ipswichian interglacial (=Eemian, approx. MIS 5e) including freshwater, then estuarine sediments. Subsequent evolution of the basin occurred during the Devensian Stage (=Weichselian, MIS 5d-2) under predominantly cold, periglacial conditions.