Graduation date: 1966

Morphogenesis in walled organisms represents a highly controlled process by which the variability of shapes arises through changes in the structure and mechanics of the cell wall. Despite taking different evolutionary paths, land plants and some brown algae exhibit great developmental and morphological similarities. In two brown algal model systems: the Sargassum muticum apex and the Fucus serratus embryo, I have used a combination of imaging techniques, growth analyses, surgical and pharmacological treatments, as well as molecular, biochemical and mechanical approaches to characterise the growth patterns and the cell wall contribution to shape change. To understand how the adult algal body is formed, I examined the branching strategy (phyllotaxis) in S. muticum. My results suggest that in S. muticum the spiral phyllotactic pattern and the apical cell division pattern are not linked. The phytohormone auxin and the biochemical changes of the cell wall do not seem to be correlated with the bud outgrowth, contrary to observations in plants. In summary, these results suggest Sargassum convergently developed a distinct growth mechanism with similar shape outcome as observed in plants. This dissertation is one of the first attempts to explore cell wall mechanics in brown algal development and its correlation with underlying cell wall biochemistry utilising the Fucus embryo as a known system. The results suggest a correlation between the wall mechanics and alginate biochemistry with the growing and non-growing regions of the embryo. In addition, altering cell wall deposition or composition has a strong effect on embryo rhizoid elongation and is, in certain cases, accompanied by significant increase in cell wall stiffness and reduction of alginate epitopes. Furthermore, preliminary results exploring transcriptomic changes during development indicate differential expression of particular alginate biosynthesis enzymes (mannuronan C5 epimerases) during development, suggesting alginate conformational modifications might be stage specific. These results contribute to the current knowledge addressing the importance of cell walls in brown algal development using novel tools and approaches. Understanding developmental processes in brown algae will provide a better insight how similar morphogenetic traits are established using different body-building mechanisms.

Philosophiae Doctor - PhD / Recently, a great deal of interest has developed towards the isolation of bioactive compounds from marine sources due to their numerous health benefits. Furthermore, marine algae are valuable sources of structurally diverse metabolites with scientifically proven therapeutic claims. The cell walls are rich in sulfated polysaccharides such as fucoidans in brown algae, carrageenans in red algae and ulvans in green algae. These sulfated polysaccharides exhibit many beneficial biological activities such as anticoagulant, antiviral, antioxidative, anticancer and immunomodulating activities. They have great potential for further development as products in cosmeceutical, pharmaceutical and nutraceutical areas. Although the mechanism of action is still not clear, their biological activities could be mainly attributed to their major secondary metabolites namely; phlorotannins, terpenoids and fucoidans. There was use of comprehensive chromatographic separations and a full analysis of isolates using one or other of the spectroscopic techniques. Antioxidant and cytotoxicity tests were perfomed in details for Ecklonia maxima. Furthermore, structural and electronic features of the phlorotannins were compared in an attempt to provide an explanation for the differences in their radical scavenging properties. In this regard, two main radical scavenging mechanisms, hydrogen atom transfer (HAT) and electron transfer (ET), were assessed in order to determine the preferred mode of radical scavenging. Fully relaxed geometry optimizations of the neutral and the radical species were performed utilizing DFT/B3LYP and DFT/UB3LYP methods respectively. In further studies, the structural and functional properties of sulfated polysaccharides from the three brown and one red seaweeds were investigated. This was through detailed analysis of chemical composition of crude and purified polysaccharides using PMP - derivatization of hydrolysed sugars, anion exchange, molecular weight determination, ion chromatography , FT-IR, NMR to methylation analysis. The work reports isolation and characterization of compounds from four algae: Ecklonia maxima, phlorotannin derivatives, namely phloroglucinol (22), eckol (23), 7-phloroeckol (24), 2-phloroeckol (25) and a sterol, 24-ethylidine cholesterol (26); Splachinidium rugosum, 24-ethylidine cholesterol ( 26), 1, 3-Dicapryloyl-2-oleoylglycerol (27), E-3,7,11,15-tetramethylhexa dec-2-en-1-ol (phytol) (2 8); Macrocystis angustifolia, 24-ethylidine cholesterol (26); a red seaweed Aeodes orbitosa, and E -3, 7, 11, 15-tetramethylhexad ec-2-en-1-ol (28) and 17-(5-Ethyl-6-methylheptan-2-yl)-10,13-dimethyl-2,3,4,7, 8,9,11,12,14,15,16,17-dodecahydro-1 H-cyclopenta[a]phenanthren-3-ol (β-sitosterol) (29). Experimental findings and theoretical predictions of phlorotannins indicated that the radical scavenging activities followed the order 22< 23 < 25 < 24. Theoretical studies further indicated the ET mechanism is more significant than the HAT mechanism due to the high BDE values. Their polysaccharide structures were tentatively shown to have a backbone of (1-3) and (1-4) linkages with sulfate groups at O-2 and O-2, 3 positions. The only red algae studied contained, 2-O-methyl-D-galactose with (1-3) and (1-4)-glycosidic linkages possessing sulfate groups at positions 2 and 6.

Summary in English. / Bibliography: pages 146-169. / The biology, population dynamics, and the environmental tolerances (temperature and light) in laboratory culture, of the endemic intertidal South African fucoid Bifurcaria brassicaeformis (Kütz) Barton have been investigated. A general description is included of the morphology and anatomy, and comparisons made with that of other species in the genus. Studies on plants collected from contrasting habitats on the lowshore and from mid-shore pools revealed that while cortical thickness of vegetative uprights were similar for both habitats, medullary diameters were generally higher, corresponding to generally thicker uprights in the low-shore. Observations on the method of zygote attachment revealed that attachment of the species resembles that of the European Bifurcaria and Halidrys siliguosa which both also display delayed rhizoidal development and initially attach by means of a mucilage secretion of the zygote wall. Observations on receptacle anatomy revealed that size of conceptacles increase from the apex to the base of a receptacle.

Brown algae are important primary producers and habitat formers in coastal environments and are believed to have evolved multicellularity independently of the other eukaryotes. The phaeoviruses that infect them form a stable lysogenic relationship with their host via genome integration, but have only been extensively studied in two genera: Ectocarpus and Feldmannia. In this study I aim to improve our understanding of the genetic diversity, host range and distribution of phaeoviruses. Sequencing and phylogenetic analysis of amplified fragments of three core phaeoviral genes (encoding major capsid protein (MCP), DNA polymerase and superfamily III helicase) of phaeovirus infected algae confirmed the suspected phaeoviral identity of viruses infecting E. fasciculatus, F. simplex, Pilayella littoralis, Myriotrichia clavaeformis and Hincksia hincksiae. Furthermore, this approach revealed multiple virus sequence variants within individual strains, and moreover that the variants formed two distinct subgroups. Subgroup A was highly conserved and observed in multiple algal genera, whereas subgroup B was much more diverse, but only found in Feldmannia species. Transcriptome sequencing of an actively infected F. irregularis strain revealed polymorphisms within key viral genes, suggesting that multiple variants were indeed active within this strain. High resolution melt curve (HRM) technology was used to develop a high throughput screening method for detecting phaeoviral MCP as a proxy for detection of phaeoviruses. This technique was also able to assign 88% of those detected to one of the subgroups, based on their differing melting temperature distributions. This was then applied to 1034 Ectocarpus isolates collected from around Europe and South America, and in accordance with previous studies of phaeoviral infection, 43-79% of strains contain virus sequence (depending on species). 17% of the isolates tested even contained sequence from both subgroups. 82 Laminariales strains, close relatives of the Ectocarpales, were also screened because they comprise commercially important kelp species but are not known to be infected by viruses. 10-17% of these tested positive for phaeoviral MCP, which when sequenced formed a separate group within the phaeoviruses. This finding could have a major impact on the kelp farming industry if the viruses are found to affect reproduction as happens in the Ectocarpales. The discovery of two subgroups is contrary to current beliefs that the phaeoviruses are a single monophyletic group, and that each species of alga has its own phaeovirus, casting doubt on the usefulness of the current convention of naming each phaeovirus after its host. It appears that the subgroup B viruses have begun to evolve away from the stable, K-selected subgroup A viruses towards a more r- type strategy with higher mutation and diversification. This study has identified potential mechanisms that may influence this shift, including mutations in a region of the DNA polymerase known to negatively affect DNA replication fidelity, combined with an active integrase and lack of a proofreading exonuclease, along with the observed infection of individuals with both phaeovirusal subgroups. The resulting mutations and recombinations could lead to the diversity observed here, and may provide a suitable model for the study of other emergent virus infections.

The aims of this study were to find endophytic brown algae in marine macroalgae from New Zealand, isolate them into culture and identify them using morphological as well as molecular markers, to study the prevalence of pigmented endophytes in a representative host-endophyte relationship, and to reveal the ultrastructure of the interface between the obligate parasite Herpodiscus durvillaeae (LINDAUER) SOUTH and its host Durvillaea antarctica (CHAMISSO) HARRIOT. Three species of pigmented endophytic Phaeophyceae were isolated from New Zealand macrophytes. They were distinguished based on morphological characters in culture, in combination with their distribution among different host species and symptoms associated with the infection of hosts. ITS1 nrDNA sequences confirmed the identity of two of the species as Laminariocolax macrocystis (PETERS) PETERS in BURKHARDT & PETERS and Microspongium tenuissimum (HAUCK) PETERS. A new genus and species, Xiphophorocolax aotearoae gen. et sp. ined., is suggested for the third group of endophytic Phaeophyceae. Three genetic varieties of L. macrocystis as well as two varieties each of M. tenuissimum and X. aotearoae were present among the isolates. L. macrocystis and X. aotearoae constitute new records for the marine flora of the New Zealand archipelago, on genus and species level. The red algal endophyte Mikrosyphar pachymeniae LINDAUER previously described from New Zealand is possibly synonymous with Microspongium tenuissimum. The prevalence of infection by Laminariocolax macrocystis was investigated in three populations of Macrocystis pyrifera along the Otago coast. Two of the populations situated inside and at the entrance of Otago Harbour showed high infection rates (average between 95 and 100%), while an offshore population was less infected (average of 35%). The phylogenetic affinities of the parasitic brown alga Herpodiscus durvillaeae, an obligate endophyte of Durvillaea antarctica (Fucales, Phaeophyceae) in New Zealand, were investigated. Analyses combined nuclear encoded ribosomal and plastid encoded RuBisCO genes. Results from parsimony, distance and likelihood methods suggest a placement of this species within the order Sphacelariales. Even though H. durvillaeae shows a reduced morphology, molecular data were supported by two morphological features characteristic for the Sphacelariales: the putative presence of apical cells and the transistory blackening of the cell wall with 'Eau de Javelle'. Ultrastructural sections showed evidence for a symplastic contact between the cells of the parasite H. durvillaeae and its host D. antarctica. Within the host cortex, parasite cells attack the fields of plasmodesmata connecting host cells. In these areas, parasite cells squeeze between the host cells and form secondary plasmodesmata connecting the primary plasmodesmata of the host cells with the cytoplasma of the parasite cell. Moreover, despite being described as lacking pigments, H. durvillaeae possesses a rbcL gene, and its plastids show red autofluorescence in UV light, suggesting the presence of a possibly reduced, but functional photosynthetic apparatus. Vestigial walls between developing spores in the 'secondary unilocular sporangia' of H. durvillaeae confirm the identity of these sporangia as plurilocular gametangia, derived from reduced gametophytes which were entirely transformed into gametangia.

Durvillaea, a kelp genus occurring only in the Southern Hemisphere, presents an ideal system for studies of marine connectivity and postglacial recolonisation. Durvillaea contains five currently-recognised species, four of which are non-buoyant. Whereas all non-buoyant species are restricted to the south-western Pacific, the sole buoyant species (D. antarctica) has a far wider, circumpolar distribution, strongly suggesting that long-distance dispersal in D. antarctica is achieved by rafting. This contrast in predicted dispersal ability among Durvillaea species provides an opportunity for natural phylogeographic comparisons, thereby assessing the effectiveness of rafting as a long-distance dispersal mechanism. Additionally, the inability of D. antarctica to survive in ice-affected areas, combined with its broad distribution, make it an ideal candidate for studies of postglacial recolonisation. Phylogenetic and biogeographic relationships within Durvillaea were here assessed using sequence data from mitochondrial (COI), chloroplast (rbcL) and nuclear (18S) DNA. Genetic data were obtained from more than 500 specimens, including representatives from across the geographic range of each recognised species of Durvillaea. Mitochondrial data for Durvillaea were found to be highly phylogenetically informative, with 117 variable sites observed over a 629 bp fragment of COI. Chloroplast and nuclear markers, on the other hand, showed less variation than COI, but nonetheless contributed useful phylogenetic information. Phylogenetic analyses were performed using both Maximum Likelihood and Bayesian approaches. Contrasting patterns of genetic diversity were observed across the range of D. potatorum in Australia, with genetic homogeneity throughout western sites versus relatively high levels of diversity in eastern populations. Based on these results, I hypothesise that D. potatorum recolonised much of the western part of its range postglacially, perhaps being entirely eliminated from western Tasmania during the last glacial period by altered oceanographic systems. Additionally, 'western' and 'eastern' D. potatorum haplotypes formed deeply-divergent clades, likely reflecting geographic isolation on either side of the Bassian Isthmus during Pleistocene marine regressions. Substantial genetic diversity was observed across the range of the circumpolar species D. antarctica. Within New Zealand, phylogenetic and morphological analyses of D. antarctica indicate that two morphotypes ('cape' and 'thonged' forms) likely represent reproductively isolated species, with the 'cape' lineage apparently restricted to southern New Zealand. Whereas the 'cape' lineage showed little genetic variation throughout its range, the 'thonged' lineage exhibited marked phylogeographic structure, with high genetic diversity and a clear north - south genetic disjunction delineated by the Canterbury Bight. On a broader, circumpolar scale, D. antarctica showed contrasting patterns of genetic diversity, with high levels of variation in low-latitude regions (e.g., continental coasts of New Zealand and Chile), versus near-homogeneity at high, subantarctic latitudes. These phylogeographic contrasts strongly suggest that D. antarctica recolonised much of the subantarctic region only recently, most plausibly following extirpation by ice scour at the Last Glacial Maximum (LGM). Locations of putative recolonised islands relative to 'refugial' areas indicate that Antarctic sea ice was likely more extensive at the LGM than previous studies have suggested. Latitudinal contrasts in genetic diversity were also observed among Chilean populations of D. antarctica, with a single mitochondrial haplotype detected throughout Chilean Patagonia versus high diversity in central Chile (32�-42�S). The Patagonian populations appear to have recolonised the region postglacially, following recession of the Patagonian Ice Sheet after the LGM. These populations show transoceanic ancestry, with a closer relationship to populations in the subantarctic and southern New Zealand regions than to those in central Chile. Substantial phylogeographic structure was evident across small spatial scales in central Chile, and the correspondence of major genetic disjunctions among central Chilean sites with the presence of long stretches of unsuitable substrate (beaches) strongly suggests that habitat discontinuity drives genetic isolation in this dispersive species. Broad-scale molecular phylogenetic analyses indicate that the current taxonomy of Durvillaea species requires substantial revision. Previously-recognised 'morphotypes' of Durvillaea (of D. potatorum in Australia, and D. antarctica in New Zealand), for example, were here found to be genetically distinct, likely representing reproductively-isolated species. This phylogeographic research on Durvillaea kelp sheds new light on the historical impacts of climate change on Southern Hemisphere marine environments, and on the processes driving evolution in a marine macroalga.

Durvillaea, a kelp genus occurring only in the Southern Hemisphere, presents an ideal system for studies of marine connectivity and postglacial recolonisation. Durvillaea contains five currently-recognised species, four of which are non-buoyant. Whereas all non-buoyant species are restricted to the south-western Pacific, the sole buoyant species (D. antarctica) has a far wider, circumpolar distribution, strongly suggesting that long-distance dispersal in D. antarctica is achieved by rafting. This contrast in predicted dispersal ability among Durvillaea species provides an opportunity for natural phylogeographic comparisons, thereby assessing the effectiveness of rafting as a long-distance dispersal mechanism. Additionally, the inability of D. antarctica to survive in ice-affected areas, combined with its broad distribution, make it an ideal candidate for studies of postglacial recolonisation. Phylogenetic and biogeographic relationships within Durvillaea were here assessed using sequence data from mitochondrial (COI), chloroplast (rbcL) and nuclear (18S) DNA. Genetic data were obtained from more than 500 specimens, including representatives from across the geographic range of each recognised species of Durvillaea. Mitochondrial data for Durvillaea were found to be highly phylogenetically informative, with 117 variable sites observed over a 629 bp fragment of COI. Chloroplast and nuclear markers, on the other hand, showed less variation than COI, but nonetheless contributed useful phylogenetic information. Phylogenetic analyses were performed using both Maximum Likelihood and Bayesian approaches. Contrasting patterns of genetic diversity were observed across the range of D. potatorum in Australia, with genetic homogeneity throughout western sites versus relatively high levels of diversity in eastern populations. Based on these results, I hypothesise that D. potatorum recolonised much of the western part of its range postglacially, perhaps being entirely eliminated from western Tasmania during the last glacial period by altered oceanographic systems. Additionally, 'western' and 'eastern' D. potatorum haplotypes formed deeply-divergent clades, likely reflecting geographic isolation on either side of the Bassian Isthmus during Pleistocene marine regressions. Substantial genetic diversity was observed across the range of the circumpolar species D. antarctica. Within New Zealand, phylogenetic and morphological analyses of D. antarctica indicate that two morphotypes ('cape' and 'thonged' forms) likely represent reproductively isolated species, with the 'cape' lineage apparently restricted to southern New Zealand. Whereas the 'cape' lineage showed little genetic variation throughout its range, the 'thonged' lineage exhibited marked phylogeographic structure, with high genetic diversity and a clear north - south genetic disjunction delineated by the Canterbury Bight. On a broader, circumpolar scale, D. antarctica showed contrasting patterns of genetic diversity, with high levels of variation in low-latitude regions (e.g., continental coasts of New Zealand and Chile), versus near-homogeneity at high, subantarctic latitudes. These phylogeographic contrasts strongly suggest that D. antarctica recolonised much of the subantarctic region only recently, most plausibly following extirpation by ice scour at the Last Glacial Maximum (LGM). Locations of putative recolonised islands relative to 'refugial' areas indicate that Antarctic sea ice was likely more extensive at the LGM than previous studies have suggested. Latitudinal contrasts in genetic diversity were also observed among Chilean populations of D. antarctica, with a single mitochondrial haplotype detected throughout Chilean Patagonia versus high diversity in central Chile (32�-42�S). The Patagonian populations appear to have recolonised the region postglacially, following recession of the Patagonian Ice Sheet after the LGM. These populations show transoceanic ancestry, with a closer relationship to populations in the subantarctic and southern New Zealand regions than to those in central Chile. Substantial phylogeographic structure was evident across small spatial scales in central Chile, and the correspondence of major genetic disjunctions among central Chilean sites with the presence of long stretches of unsuitable substrate (beaches) strongly suggests that habitat discontinuity drives genetic isolation in this dispersive species. Broad-scale molecular phylogenetic analyses indicate that the current taxonomy of Durvillaea species requires substantial revision. Previously-recognised 'morphotypes' of Durvillaea (of D. potatorum in Australia, and D. antarctica in New Zealand), for example, were here found to be genetically distinct, likely representing reproductively-isolated species. This phylogeographic research on Durvillaea kelp sheds new light on the historical impacts of climate change on Southern Hemisphere marine environments, and on the processes driving evolution in a marine macroalga.

Kelpak®, a seaweed concentrate (SWC) prepared from the brown seaweed Ecklonia maxima (Osbeck) Papenfuss, improves overall plant mass and fruit yield in a variety of crops. The main active principals isolated from Kelpak® are cytokinins and auxins. Although these compounds are partly responsible for the growth promoting effect observed with Kelpak® application, they do not fully account for the complete effect of Kelpak® treatment. For this reason the focus has turned to polyamines (PAs) which are found in all cells of plants, animals and microorganisms, including eukaryotic algae. Polyamines also have growth promoting effects in plants. A study was carried out to investigate the PA levels in E. maxima and Kelpak® through a biennial cycle and to investigate if the PAs present in Kelpak® may have an effect on root growth, alleviating nutrient deficiency and the transport and accumulation of PAs in plants. To determine the amount of PA in the stipes, fronds and SWC prepared from E. maxima, samples were collected monthly over a two-year period (June 2009-June 2011). Extracts were benzoylated and quantified using a Varian HPLC. Putrescine concentrations ranged from 15.98-54.46 μg.g⁻¹, 6.01-40.46 μg.g⁻¹ and 50.66-220.49 μg.g⁻¹ DW in the stipe, fronds and SWC, respectively. Spermine concentrations ranged from 1.02-35.44 μg.g⁻¹, 1.05-26.92 μg.g⁻¹ and 7.28-118.52 μg.g⁻¹ DW in the stipe, fronds and SWC, respectively. Spermidine concentrations fell below the detection threshold. This is the first report of PAs being detected in a SWC. The seasonal pattern established for the stipe, frond and SWC followed the same trend over a biennial cycle. Polyamines accumulated in the seaweed tissue during periods of active growth and as a stress response elicited by rough wave action. This PA trend was similar to the cytokinin trend reported by MOONEY and VAN STADEN (1984b) for Sargassum heterophyllum which suggests that PAs play an important role in the hormone cascade during active growth. Routine monthly screening of Kelpak® carried out in the Research Centre for Plant Growth and Development indicated that Kelpak® consistently resulted in more rooting in the mung bean bioassay than the IBA control. The potential root promoting effect of PAs were investigated. Individually applied PAs did not increase rooting in the mung bean bioassay, but a synergistic relationship was observed between Put (10⁻³ M) and IBA (10⁻⁴ M). When applied together, rooting increased significantly above Put (10⁻³ M) and IBA (10⁻⁴ M) applied separately. The Put-auxin combination produced a similar number of roots to those treated with Kelpak®. It is possible that the PAs present in Kelpak® have a synergistic effect with auxins present in Kelpak® to promote root development and growth. Several physiological effects of Kelpak® and PAs on plant growth were investigated in a series of pot trials. Kelpak® significantly improved the growth of P- and K-deficient okra seedlings and masked the detrimental effects exerted by P- and K-deficiency. The application of PAs (10⁻⁴ M) significantly improved the seedling vigour index (SVI) of okra seedlings subjected to N-deficiency. The statistical difference was attributed to the N-containing growth regulators and polyamines being degraded and metabolized by the okra seedlings. Polyamine application did not alleviate P- and K-deficiency but increased root growth significantly in seedlings receiving an adequate supply of nutrients. It is likely that the additional PAs supported auxin-mediated root growth. A pot trial with okra plants was conducted to establish if the PAs in Kelpak®, applied as a soil drench or foliar application, are absorbed and translocated in a plant. Plants were also treated with Put, Spm, Spd to establish if PAs can be absorbed and translocated. Once the fruit had matured, plants were harvested and the endogenous PA content quantified by HPLC in the roots, stems and fruits. Applying PAs as a soil drench was not as effective as a foliar spray at increasing the PA content in the different plant parts. Kelpak® treatment (0.4%) did not contribute more PAs in any plant part. Spermidine concentrations were higher, in the various plant parts, than Put or Spm, irrespective of the mode of application. The application of Put, Spd and Spm increased Spd concentrations in the roots. Considering that Spd is the main PA produced in the roots and that exogenously applied PAs are readily converted to Spd, it seems evident that Spd is the preferred PA for long-distance transport in plants. The cytokinins and auxins in Kelpak® play an important role in stimulating growth in plants. It is, however, the totality of different compounds in Kelpak® that gives it its unique growth stimulating ability. Polyamines, occurring within the seaweed contribute to this activity, having an active role in root production and thus increased plant growth. / Thesis (M.Sc.)-University of KwaZulu-Natal, Pietermaritzburg, 2011.

Un premier pan de ma thèse concerne deux différents mécanismes de sauvetage du Y par le X. Premièrement, j'ai participé à une controverse sur la compensation de dosage chez les mammifères. Une hypothèse avait été proposée dans les années 60 par Susumo Ohno, proposant un mécanisme de compensation en deux temps. Chez les mâles, la perte de nombreux gènes sur le Y entraîne un déséquilibre de dosage car ces gènes qui étaient précédemment présents en deux copies sont devenus unicopie, soit une division d'expression par deux. Selon l'hypothèse d'Ohno, chez les mammifères en réponse à cela le X aurait doublé son expression, mais dans les deux sexes menant ainsi à une expression trop élevée chez les femelles. Ce deuxième problème de dosage aurait alors été résolu par la mise en place d'une inactivation aléatoire de l'un des deux X chez les femelles. Tandis que la deuxième partie de l'hypothèse d'Ohno, l'inactivation du X, a été très étudiée, la première partie est restée spéculative jusqu'aux années 2000. En étudiant des données d'expression du X humain j'ai pu montrer, de manière concomitante avec d'autres auteurs, que la première partie de l'hypothèse d'Ohno n'est pas totalement vraie car seule une partie des gènes sont sur-exprimés. J'ai ensuite participé à l'écriture d'une revue visant à donner une explication alternative à la compensation de dosage pour l'évolution de l'inactivation du X chez les femelles mammifères. Deuxièmement, j'ai étudié la présence de conversion génique X-Y dans plusieurs gènes, au sein de nombreuses espèces de primates. Mes travaux me mènent à discuter le fait que ce type d'évènement soit effectivement favorisé par la sélection. Je pose l'hypothèse que ces conversions géniques ont été maintenues de manière neutre. Ces deux travaux ne vont pas dans le sens d'un chromosome X sauvant le Y avec beaucoup de zèle. Dans un dernier temps, m'éloignant des espèces modèles, j'ai étudié les chromosomes sexuels particuliers d'une algue brune : Ectocarpus siliculosus. Cela m'a permis de vérifier si le scénario évolutif actuel des chromosomes sexuels est toujours valide dans un groupe d'eucaryotes séparé des animaux depuis plus d'un milliard d'années / The first part of my thesis concerns two different mechanisms of the Y being rescued by the X. Firstly, I contributed to a controversy on mammalian dosage compensation. During the 60s Susumo Ohno hypothesized a two-step dosage compensation mechanism. In males, the high loss of Y-linked genes led to a dosage imbalance: these genes were previously present in two allelic copies and became unicopy, meaning that their expression has been halved. According to Ohno’s hypothesis, in response to this imbalance the mammalian X would have doubled its expression in the two sexes, resulting in a to high expression in females. This second dosage imbalance would have been resolved by the random inactivation of one of the two Xs in females. Whereas the second part of Ohno’s hypothesis, the X-chromosome inactivation, has been well studied, the first part remained speculative until the 2000s. I studied human X-linked expression data and was able to show, concomitantly with other authors, that the first part of Ohno’s hypothesis is not totally true as only some of the X-linked genes are hyperexpressed. I later participated in the writing of a review aiming to give an alternative hypothesis for the evolution of X-chromosome inactivation in mammalian females than dosage compensation. Secondly, I studied signatures of X-Y gene conversion in several genes within numerous primate species. Myresults led me to discuss if these events were indeed selected for. I hypothesize that these gene conversion events occurred in a neutral manner. These two different studies suggest that the X chromosome may not be as much a help for the Y as has been suggested. Lastly, moving away from model species, I studied the peculiar sex chromosomes of a brown alga: Ectocarpus siliculosus. This work allowed me to test if the current hypotheses on sex chromosome evolution still hold in a eukaryotic group that diverged from animals more than one billion years ago