Theoretical aspects of the evolution of reproductive parasites.

Reproductive parasites are maternally inherited endosymbionts that manipulate the reproduction of their hosts in a way that enhances the transmission of the parasites, but is deleterious to the hosts. In the present thesis, I try to resolve some ques tions concerning the evolution of reproductive parasites and their hosts by means of theoretical modelling, using a variety of approaches including recurrence equations, optimisation, and stochastic modelling. I first study the question: 'Can male-killing bacteria and meiotic drive elements influence each others' spread and equilibrium frequency in a population' and demonstrate they can. Following this, I examine two questions with respect to host evolution. First, can male-killing or Cl-inducing bacteria facilitate the evolution of haplodiploidy I conclude that past work on this has been overoptimistic, but that the process is possible. Second, how does the pres ence of male-killing bacteria affect basic population genetic processes, in particular the interplay between natural selection and random genetic drift I demonstrate that the host population behaves approximately as if only uninfected individuals existed. I then examine two questions in relation to endosymbiont biology. First, what is the impact of mating systems and segregation on the evolution of new Cl-types I propose outbreeding systems as a context where new CI-types can evolve. Second, how can we expect selection to act on two bacterial strains of reproductive parasites in doubly infected hosts with respect to their density Finally, I examine how we expect the incidence of reproductive parasites to vary in time within a host clade, and how the phylogenetic history of the host clade affects the pattern of spread and expected incidence of the parasites. I conclude my thesis with a general overview of what is known about the evolution of reproductive parasites both empirically and theoretically, and outline some promising future avenues of research

Links between topography, wind, deflation, lakes and dust: The case of the Bodélé Depression, Chad

The Bodélé Depression, Chad is the planet's largest single source of dust. Deflation from the Bodélé could be seen as a simple coincidence of two key prerequisites: strong surface winds and a large source of suitable sediment. But here we hypothesise that long term links between topography, winds, deflation and dust ensure the maintenance of the dust source such that these two apparently coincidental key ingredients are connected by land-atmosphere processes with topography acting as the overall controlling agent. We use a variety of observational and numerical techniques, including a regional climate model, to show that: 1) contemporary deflation from the Bodélé is delineated by topography and a surface wind stress maximum; 2) the Tibesti and Ennedi mountains play a key role in the generation of the erosive winds in the form of the Bodélé Low Level Jet (LLJ); 3) enhanced deflation from a stronger Bodélé LLJ during drier phases, for example, the Last Glacial Maximum, was probably sufficient to create the shallow lake in which diatoms lived during wetter phases, such as the Holocene pluvial. Winds may therefore have helped to create the depression in which erodible diatom material accumulated. Instead of a simple coincidence of nature, dust from the world's largest source may result from the operation of long term processes on paleo timescales which have led to ideal conditions for dust generation in the world's largest dust source. Similar processes plausibly operate in other dust hotspots in topographic depressions

A detailed characterization of the Saharan dust collected during the Fennec campaign in 2011: in situ ground-based and laboratory measurements

Millions of tons of mineral dust are lifted by the wind from arid surfaces and transported around the globe every year. The physical and chemical properties of the mineral dust are needed to better constrain remote sensing observations and are of fundamental importance for the understanding of dust atmospheric processes. Ground-based in situ measurements and in situ filter collection of Saharan dust were obtained during the Fennec campaign in the central Sahara in 2011. This paper presents results of the absorption and scattering coefficients, and hence single scattering albedo (SSA), of the Saharan dust measured in real time during the last period of the campaign and subsequent laboratory analysis of the dust samples collected in two supersites, SS1 and SS2, in Algeria and in Mauritania, respectively. The samples were taken to the laboratory, where their size and aspect ratio distributions, mean chemical composition, spectral mass absorption efficiency, and spectral imaginary refractive index were obtained from the ultraviolet (UV) to the near-infrared (NIR) wavelengths. At SS1 in Algeria, the time series of the scattering coefficients during the period of the campaign show dust events exceeding 3500 Mm−1, and a relatively high mean SSA of 0.995 at 670 nm was observed at this site. The laboratory results show for the fine particle size distributions (particles diameter  < 5µm and mode diameter at 2–3 µm) in both sites a spectral dependence of the imaginary part of the refractive index Im(m) with a bow-like shape, with increased absorption in UV as well as in the shortwave infrared. The same signature was not observed, however, in the mixed particle size distribution (particle diameter  < 10 µm and mode diameter at 4 µm) in Algeria. Im(m) was found to range from 0.011 to 0.001i for dust collected in Algeria and 0.008 to 0.002i for dust collected in Mauritania over the wavelength range of 350–2500 nm. Differences in the mean elemental composition of the dust collected in the supersites in Algeria and in Mauritania and between fine and mixed particle size distributions were observed from EDXRF measurements, although those differences cannot be used to explain the optical properties variability between the samples. Finally, particles with low-density typically larger than 10 µm in diameter were found in some of the samples collected at the supersite in Mauritania, but these low-density particles were not observed in Algeria

Detection and analysis of Lhù'ààn Mân' (Kluane Lake) dust plumes using passive and active ground-based remote sensing supported by physical surface measurements

There is growing recognition that high-latitude dust (HLD), originating from local drainage-basin flows, is the dominant source for certain important phenomena such as particle deposition on snow/ice. The analysis of such local plumes (including a better exploitation of remote sensing data) has been targeted as a key aerosol issue by the HLD community. The sub-Arctic Lhù'ààn Mân' (Kluane Lake) region in the Canadian Yukon is subject to regular drainage-basin, wind-induced dust plumes. This dust emission site is one of many current and potential proglacial dust sources in the Canadian north. In situ ground-based measurements are, due to constraints in accessing these types of regions, rare. Ground- and satellite-based remote sensing accordingly play an important role in helping to characterize local dust sources in the Arctic and sub-Arctic. We compared ground-based passive and active remote sensing springtime (May 2019) retrievals with microphysical surface-based measurements in the Lhù'ààn Mân' region in order to better understand the potential for ground- and satellite-based remote sensing of HLD plumes. This included correlation analyses between ground-based coarse mode (CM) aerosol optical depth (AOD) retrievals from AERONET AOD spectra, CM AODs derived from co-located Doppler lidar profiles, and OPS (optical particle sizer) surface measurements of CM particle-volume concentration (vc(0)). An automated dust classification scheme was developed to objectively identify local dust events. The classification process helped distinguish lidar-derived CM AODs which covaried with vdust(0) (during recognized dust events) and those that varied at the same columnar scale as AERONET-derived CM AOD (and thus could be remotely sensed). False positive cloud events for dust-induced, high-frequency variations in lidar-derived CM AODs in cloudless atmospheres indicated that the AERONET cloud-screening process was rejecting CM dust AODs. The persistence of a positive lidar ratio bias in comparing the CIMEL/lidar-derived value with a prescribed value obtained from OPS-derived particle sizes coupled with dust-speciation-derived refractive indices led to the suggestion that the prescribed value could be increased to optically derived values of 20 sr by the presence of optically significant dust particles at an effective radius of 11–12 µm. Bimodal CM PSDs (see Appendix B for a glossary) from full-fledged AERONET inversions (the combination of AOD spectra and almucantar radiances) also showed CM peaks at ∼ 1.3 and 5–6.6 µm radius: this, we argue, was associated with springtime Asian dust and Lhù'ààn Mân' dust, respectively. Correlations between the CIMEL-derived fine mode (FM) AOD and FM OPS-derived particle-volume concentrations suggest that remote sensing techniques can be employed to monitor FM dust (which is arguably a better indicator of the long-distance transport of HLD).</p

Advances in understanding mineral dust and boundary layer processes over the Sahara from Fennec aircraft observations

International audienceThe Fennec climate program aims to improve understanding of the Saharan climate system through a synergy of observations and modelling. We present a description of the Fennec airborne observations during 2011 and 2012 over the remote Sahara (Mauritania and Mali) and the advances in the understanding of mineral dust and boundary layer processes they have provided. Aircraft instrumentation aboard the UK FAAM BAe146 and French SAFIRE Falcon 20 is described, with specific focus on instrumentation specially developed and relevant to Saharan meteorology and dust. Flight locations, aims and associated meteorology are described. Examples and applications of aircraft measurements from the Fennec flights are presented, highlighting new scientific results delivered using a synergy of different instruments and aircraft. These include: (1) the first airborne measurement of dust particles sized up to 300 microns and associated dust fluxes in the Saharan atmospheric boundary layer (SABL), (2) dust uplift from the breakdown of the nocturnal low-level jet before becoming visible in SEVIRI satellite imagery, (3) vertical profiles of the unique vertical structure of turbulent fluxes in the SABL, (4) in-situ observations of processes in SABL clouds showing dust acting as CCN and IN at −15 °C, (5) dual-aircraft observations of the SABL dynamics, thermodynamics and composition in the Saharan heat low region (SHL), (6) airborne observations of a dust storm associated with a cold-pool (haboob) issued from deep convection over the Atlas, (7) the first airborne chemical composition measurements of dust in the SHL region with differing composition, sources (determined using Lagrangian backward trajectory calculations) and absorption properties between 2011 and 2012, (8) coincident ozone and dust surface area measurements suggest coarser particles provide a route for ozone depletion, (9) discrepancies between airborne coarse mode size distributions and AERONET sunphotometer retrievals under light dust loadings. These results provide insights into boundary layer and dust processes in the SHL region – a region of substantial global climatic importance

Quantifying the response of the ORAC aerosol optical depth retrieval for MSG SEVIRI to aerosol model assumptions

We test the response of the Oxford-RAL Aerosol and Cloud (ORAC) retrieval algorithm for MSG SEVIRI to changes in the aerosol properties used in the dust aerosol model, using data from the Dust Outflow and Deposition to the Ocean (DODO) flight campaign in August 2006. We find that using the observed DODO free tropospheric aerosol size distribution and refractive index increases simulated top of the atmosphere radiance at 0.55 µm assuming a fixed erosol optical depth of 0.5 by 10–15 %, reaching a maximum difference at low solar zenith angles. We test the sensitivity of the retrieval to the vertical distribution f the aerosol and find that this is unimportant in determining simulated radiance at 0.55 µm. We also test the ability of the ORAC retrieval when used to produce the GlobAerosol dataset to correctly identify continental aerosol outflow from the African continent and we find that it poorly constrains aerosol speciation. We develop spatially and temporally resolved prior distributions of aerosols to inform the retrieval which incorporates five aerosol models: desert dust, maritime, biomass burning, urban and continental. We use a Saharan Dust Index and the GEOS-Chem chemistry transport model to describe dust and biomass burning aerosol outflow, and compare AOD using our speciation against the GlobAerosol retrieval during January and July 2006. We find AOD discrepancies of 0.2–1 over regions of intense biomass burning outflow, where AOD from our aerosol speciation and GlobAerosol speciation can differ by as much as 50 - 70 %

Wolbachia-Induced Unidirectional Cytoplasmic Incompatibility and Speciation: Mainland-Island Model

Bacteria of the genus Wolbachia are among the most common endosymbionts in the world. In many insect species these bacteria induce a sperm-egg incompatibility between the gametes of infected males and uninfected females, commonly called unidirectional cytoplasmic incompatibility (CI). It is generally believed that unidirectional CI cannot promote speciation in hosts because infection differences between populations will be unstable and subsequent gene flow will eliminate genetic differences between diverging populations. In the present study we investigate this question theoretically in a mainland-island model with migration from mainland to island. Our analysis shows that (a) the infection polymorphism is stable below a critical migration rate, (b) an (initially) uninfected “island” can better maintain divergence at a selected locus (e.g. can adapt locally) in the presence of CI, and (c) unidirectional CI selects for premating isolation in (initially) uninfected island populations if they receive migration from a Wolbachia-infected mainland. Interestingly, premating isolation is most likely to evolve if levels of incompatibility are intermediate and if either the infection causes fecundity reductions or Wolbachia transmission is incomplete. This is because under these circumstances an infection pattern with an infected mainland and a mostly uninfected island can persist in the face of comparably high migration. We present analytical results for all three findings: (a) a lower estimation of the critical migration rate in the presence of local adaptation, (b) an analytical approximation for the gene flow reduction caused by unidirectional CI, and (c) a heuristic formula describing the invasion success of mutants at a mate preference locus. These findings generally suggest that Wolbachia-induced unidirectional CI can be a factor in divergence and speciation of hosts

CpG-island methylation of the ER promoter in colorectal cancer: analysis of micrometastases in lymph nodes from UICC stage I and II patients

Patients with UICC stage II colorectal cancer (CRC) have a risk of approximately 20% to develop disease recurrence after tumour resection. The presence and significance of micrometastases for locoregional recurrence in these patients lacking histopathological lymph node involvement on routine stained HE sections is undefined. Oestrogen receptor (ER) promoter methylation has earlier been identified in CRC. Therefore, we evaluated the methylation status of the ER promoter in lymph nodes from 49 patients with CRC UICC stage I and II as a molecular marker of micrometastases and predictor of local recurrence. DNA from 574 paraffin-embedded lymph nodes was isolated and treated with bisulphite. For the detection of methylated ER promoter sequences, quantitative real-time methylation-specific PCR was used. Of the 49 patients tested, 15 (31%) had ER methylation-positive lymph nodes. Thirteen of those (86%) remained disease free and two (14%) developed local recurrence. In the resected lymph nodes of 34 of the 49 patients (69%), no ER promoter methylation could be detected and none of these patients experienced a local relapse. The methylation status of the ER promoter in lymph nodes of UICC stage I and II CRC patients may be a useful marker for the identification of patients at a high risk for local recurrence

Sex Determination:Why So Many Ways of Doing It?

Sexual reproduction is an ancient feature of life on earth, and the familiar X and Y chromosomes in humans and other model species have led to the impression that sex determination mechanisms are old and conserved. In fact, males and females are determined by diverse mechanisms that evolve rapidly in many taxa. Yet this diversity in primary sex-determining signals is coupled with conserved molecular pathways that trigger male or female development. Conflicting selection on different parts of the genome and on the two sexes may drive many of these transitions, but few systems with rapid turnover of sex determination mechanisms have been rigorously studied. Here we survey our current understanding of how and why sex determination evolves in animals and plants and identify important gaps in our knowledge that present exciting research opportunities to characterize the evolutionary forces and molecular pathways underlying the evolution of sex determination